United States Department of DDifferencesifferences BBetweenetween Agriculture Forest Service SStandingtanding aandnd DDownedowned Northern Research Station DDeadead TTreeree WWoodood DDensityensity Research Paper NRS-15 RReductioneduction FFactors:actors: A CComparisonomparison AAcrosscross DDecayecay CClasseslasses aandnd TTreeree SSpeciespecies MMarkark E.E. HarmonHarmon CChristopherhristopher WW.. WWoodalloodall BBeckyecky FasthFasth JJayay SextonSexton MMishaisha YatkovYatkov Abstract Woody detritus or dead wood is an important part of forest ecosystems and has now become a routine facet of forest monitoring and inventory. Biomass and carbon estimates of dead wood depend on knowledge of species- and decay class- specifi c density or density reduction factors. While some progress has been made in determining these parameters for dead and downed trees (DD), there are very few estimates of these key parameters for standing dead trees (SD). In this study we evaluated indicators of decay to relate subjective SD and DD decay classifi cations then compared SD and DD density and density reduction factors by decay class for a total of 19 tree species at nine sites in the United States and Russia. A set of preliminary decay reduction factors for SD trees by species and decay class was developed for tree species inventoried by the U.S. Department of Agriculture’s Forest Inventory and Analysis (FIA) program. Results indicate that SD density declined with decay class for all examined species. For six of the examined species, SD tree density could be assumed equal to that of DD tree density. The most common situation (13 species) was for SD density to be higher than for DD density. The likely cause of these differences was the drier microenvironment of SD which slows decomposition relative to that of DD. By applying these results, a new set of SD density reduction factors was developed for 260 species inventoried by FIA in forests of the United States. Comparison of biomass estimates using this study’s proposed SD density reduction factors with existing SD volume estimates based solely on undecayed wood density indicated a possible biomass overestimate of 16.5 percent for Minnesota. Given the size of the potential biases involved and the uncertainty associated with decay reduction factors developed in this study, additional work to develop a multispecies SD decay class system and further empirical sampling of SD tree density is recommended.

Cover Photo Standing dead tree on the shores of Octopus Lake, Boundary Waters Canoe Area Wilderness, Superior National Forest, Minnesota, USA. Christopher W. Woodall, U.S. Forest Service.

Manuscript received for publication 19 April 2011

Published by: For additional copies: U.S. FOREST SERVICE U.S. Forest Service 11 CAMPUS BLVD SUITE 200 Publications Distribution NEWTOWN SQUARE PA 19073 359 Main Road Delaware, OH 43015-8640 Fax: (740)368-0152 August 2011 Email: [email protected]

Visit our homepage at: http://www.nrs.fs.fed.us/ INTRODUCTION Woody detritus or dead wood is an important part of forest ecosystems, providing habitats and food sources, a store of carbon, nutrients, and water, as well as infl uencing geomorphic processes (Franklin et al. 1987, Harmon et al. 1986, Triska and Cromack 1980). In particular, standing dead trees provide structural complexity and enhance biodiversity in many forest ecosystems (Jonsell et al. 1998, Kruys et al. 1999, Nilsson et al. 2001). Th erefore, dead wood is increasingly being included in ecological studies and environmental assessments (Woodall et al. 2009), with the number of publications on this subject rapidly growing. Both downed (DD) and standing dead (SD) wood store carbon and are important components of forest carbon dynamics, Th e Authors but also release carbon via decomposition and combustion in fi res. MARK E. HARMON is a Richardson However, given the contrasting microenvironments that SD and DD Chair and professor in forest science with the trees inhabit, their decay dynamics and attributes may vary (Aakala Department of Forest Science, Oregon State et al. 2008, Aakala 2010, Lee 1998, Vanderwel et al. 2006). Accurate University, Corvallis, OR. estimates of standing dead tree carbon stocks are important for monitoring carbon fl ux, for characterizing processes such as respiration CHRISTOPHER W. WOODALL is research and combustion losses, and for validating simulation models that are forester with the U.S. Forest Service, Northern used to project how carbon dynamics will change in the future. Research Station’s Forest Inventory and Analysis unit, St, Paul MN. It is diffi cult to develop accurate estimates of dead wood biomass for several reasons. First, until recently there have been few systematic, BECKY FASTH is a senior research assistant regional-scale dead wood inventories, resulting in highly uncertain with the Department of Forest Science, Oregon dead wood population estimates (Harmon et al. 2001). However, State University, Corvallis, OR. regional-scale inventories of dead wood are becoming more common (Woodall et al. 2009). For example, the Forest Inventory and Analysis JAY SEXTON is a senior research assistant (FIA) program of the U.S. Forest Service now conducts a national with the Department of Forest Science, Oregon inventory of woody material including fi ne woody detritus (FWD) State University, Corvallis, OR. and DD (for more details see Woodall and Monleon 2008). Second, given the size and actively decomposing nature of dead trees, direct MISHA YATKOV is a Ph.D. candidate with measurements of biomass during forest inventories are not possible. the Department of Forest Science, Oregon Instead, the volume of individual DD pieces is estimated from fi xed- State University, Corvallis, OR. area plot or line-intersect transect estimators (Woodall and Monleon 2008). Biomass of DD is then often estimated by multiplying volume by an estimate of density specifi c to a dead wood piece’s individual attributes (e.g., species and stage of decay). Density reduction factors may be defi ned as the ratio of the decayed density (current mass/volume) of a piece of dead wood compared to its undecayed density (initial live tree mass/volume; Miles and Smith 2009). Th e development of DD species- and decay class-specifi c density and decay reduction values for the FIA program has off ered the promise of substantially reducing the uncertainty associated with DD mass/carbon population estimates resulting from large-scale inventories (Harmon et al. 2008). Unfortunately, there are very few decay class- or species-

1 specifi c estimates of density or decay reduction factors for temperate forests of the northern hemisphere. As the SD trees. Th e lack of these estimates is likely due to the decay classifi cation system for both SD and DD trees are danger in felling snags during sampling and the historical qualitative (i.e., visual assessments of wood condition lack of emphasis on SD biomass/carbon estimates during and structure) and separate, the two classifi cation systems traditional forest inventories focused on estimating live need to be linked in order to propose SD to DD wood tree growing stock volume on timberlands. density relationships by decay class. First, indicators of decay were evaluated for SD and DD decay classes Several assumptions regarding wood density infl uence to ensure decay classifi cation system compatibility. biomass estimates of SD trees. In the case of the FIA Second, the density of SD and DD was compared for program, it is currently assumed that the density of the same decay classes and species. Th ese ratios were SD trees is the same as that of live trees. While this then used to estimate SD density and density reduction seems unlikely, in some situations (e.g., extremely slow factors for species that have not been sampled across the decomposition in arid environments) this assumption may United States. Finally, to initially assess the impact of be accurate. Alternately, (e.g., Smithwick et al. 2002) the wood density decay reduction factors on SD population density of SD trees for a given decay class may be assumed estimates, a SD tree inventory in Minnesota was used to be similar to that of DD wood. While this would seem to calculate state SD biomass using no decay reduction to be a more realistic assumption, it has yet to be tested factor and using factors developed in previous objectives. across a wide range of tree species and environments. Th e uncertainty associated with SD tree density is likely MATERIALS AND METHODS to have major impacts on biomass, carbon, and fuel Study Areas estimates, but without knowledge of how SD tree wood density changes through its decay process, the eff ect of To ensure our study acquired a robust estimate of SD this uncertainty cannot be quantifi ed. to DD, we sampled fi ve study sites in the United States and four in Russia representing a range of species and OBJECTIVES climates in North America and Russia (Table 1). Given the similarity between the temperate and boreal forests of Given that deadwood inventories are rapidly becoming North America and Russia, it was felt that the additional a common component of forest resource inventories statistical rigor resulting from the combination of data around the world, there is a need to refi ne the constants outweighed the loss of some regional specifi city. As study used in biomass estimation procedures for both SD and objectives focus on tree species in the United States along DD. Our objective was to determine SD species- and with greater diversity among the study sites (e.g., species decay-class specifi c density and density reduction factors composition), study site locations in the United States for forest tree species in the United States by linking were kept separate and identifi ed by unique abbreviations empirical SD data from a subset of study species with in subsequent tables, while Russian study sites were that of DD of those same species to develop constants pooled with the abbreviation “RUS.” Site descriptions for that can then be applied to other SD species. Th e broad Russian locations are based on information from Yatskov concept was to use SD:DD density relationships to utilize (2000). currently available DD density information to provide an initial approximation of SD density attributes for forest Cascade Head Experimental Forest (CHE), Oregon tree species in the United States. Cascade Head is located on the north-central coast of Oregon, 8 km north of Lincoln City. It lies entirely Harmon et al. (2008) developed a set of density within the Hebo Ranger District of the Siuslaw National reduction factors, including estimates of uncertainty, Forest. Th e site is in the Oregon Coast Range ecoregion for DD and FWD based on a thorough literature and the forests are representative of the sitka spruce- review and unpublished data. Our approach for this western hemlock (Picea sitchensis (Bong.) Carrière -Tsuga study was to sample SD and DD for 19 tree species in heterophylla (Raf.) Sarg.) and Douglas-fi r (Pseudotsuga

2 Table 1.—Site name, location, and climate description for areas sampled Site Name Climate Temperature Cascade Head EF, Oregon (CHE) Very wet Moderate Fraser EF, Colorado (FEF) Dry Cool Chapel Hill, North Carolina (NCB) Moderate Moderate Marcell EF, Minnesota (MEF) Moderate Cold Kenai Peninsula, Alaska (AK) Dry Cold St. Petersburg, Russia (RUS) Moderate Cool Krasnoyarsk, Russia (RUS) Dry Cold Irkutsk, Russia (RUS) Dry Cold Khabarovsk, Russia (RUS) Moderate Cold menziesii (Mirb.) Franco) zones of the region. Soils, project was dominated by mixed hardwoods including derived primarily from tuff aceous siltstones, are fi ne Liriodendron tulipifera L. on the moist lower slopes and textured, moderately well drained, and deep (up to mixed hardwoods with pine (Pinus taeda L. and Pinus 100+ cm). Because of the Pacifi c Ocean infl uence, CHE echinata Mill.) on the dryer mid and upper slopes. It is an has a moderate and very wet climate. Mean annual area defi ned by undulating topography, soils of poor to temperature is 10 °C with minimal seasonal variation. good quality, and a temperate climate. Soils of the study Average yearly rainfall is 2489 mm, although fog drip area include Wedowee sandy loam and Goldston slaty through the forest canopy can add 500 mm or more silt loam (Dunn 1977). Th e average annual temperature precipitation a year (Greene and Acker 1999). is 14.6 °C and the average annual precipitation is 1220 mm. Precipitation is uniformly distributed throughout Fraser Experimental Forest (FEF), Colorado the year. Th e elevation ranges from 103 to 150 m (Fasth Fraser Experimental Forest is located in the heart of the et al., in press). central Rocky Mountains, about 80 km from Denver. FEF includes subalpine forests typical of the area, with Marcell Experimental Forest (MEF), Minnesota Engelmann spruce (Picea engelmannii Parry x Engelm.) Th e Marcell Experimental Forest is an 890-ha tract of and subalpine fi r (Abies lasiocarpa (Hook.) Nutt.) land in the Chippewa National Forest, located 40 km predominating at higher elevations, and lodgepole pine north of Grand Rapids, Minnesota. Vegetation in the (Pinus contorta var. latifolia (Douglas ex Louden var. upland topography includes jack pine (Pinus banksiana latifolia Engelm. ex S. Watson)) predominating at lower Lamb.) and trembling aspen (Populus tremuloides Michx.) elevations and on drier upper slopes. Soils are generally among other species. Mineral soils of the area are derived derived from gneiss and schist and typically contain from glacial processes that occurred during and after the angular gravel and stone with very little silt or clay. Wisconsinan glaciation. Soils are mainly loamy sands Th ese soils are very permeable and can store considerable covered with fi ne sandy loam derived from eolean loess water during snowmelt. Climate varies strongly with after glacial melt. Th e climate is subhumid continental, elevation. Overall, the climate is cool and dry with long, with wide and rapid diurnal and seasonal temperature cold winters and short, cool summers. Mean annual fl uctuations. Th e average annual temperature is 3 °C temperature at the Fraser Forest headquarters is 0.5 °C and the average annual precipitation is 785 mm with 75 and mean annual precipitation over the entire FEF is 737 percent occurring in the snow-free period (mid-April to mm. (Adams et al. 2004). early November) (Adams et al. 2004).

Chapel Hill (NCB), North Carolina Kenai Peninsula (AK), Alaska Th e study was conducted at the North Carolina Botanical Th e study was conducted on the Kenai Peninsula of Garden, a 242-ha tract of oak-hickory-pine forest in Alaska on Federal lands including the Chugach National Chapel Hill, North Carolina. Th e area sampled in this Forest as well as on State owned lands. Th e maritime

3 climate of this study site supports temperate, coniferous Siberian spruce (Picea obovata Ledeb.) were common, rainforest. Th e drier and colder climate inland from and were associated with small amounts of Siberian fi r the coastal mountains supports forests dominated by (Abies siberica Ledeb.). Th e mean temperature of July is white spruce (Picea glauca (Moench) Voss), black spruce 17 °C and mean temperature of January is -20 °C with an (Picea mariana (Miller) Britt), and paper birch (Betula overall mean annual temperature of -0.6 °C and a mean papyrifera Marsh). A fertile hybrid between white spruce annual precipitation of 216 mm. and Sitka spruce (Lutz spruce, Picea x lutzii Little) is a common tree in maritime and mountainous areas of the Khabarovsk, Russia (RUS) Kenai Peninsula. Th e topography of the study area varies Th is region is part of the Far East and has a continental from lowlands to foothills. Th e lowlands of the Kenai climate that promotes the growth of a wide range of plant Peninsula are a glaciated surface with gentle relief and species, including Korean white pine (Pinus koraiensis a mean elevation of approximately 300 m. Th roughout Sieb. et Zucc.), Dahurian larch (Larix dahurica Turcz.) most of this lowland glacial till of Wisconsin age is spruce (Picea obovata and Picea ajanensis Fisch.), true fi r overlain by deep glacial lake and fl uvial sediments and is (Abies siberica), and yellow birch (Betula costata Trautv.). mantled with loess. Th e Alaska Climate Research Center Th e vegetation period lasts 174 days in lowlands and 158 (ACRC) climatic summaries indicate the study area has days in mountainous areas. Th e mean temperature of July a mean annual temperature of approximately 1.1 °C and is 17 °C, while mean temperature of January is -24 °C, annual precipitation of 630 mm (Harmon et al. 2005). with an overall annual mean of -1.9 °C. Th e mean annual precipitation is 909 mm. St. Petersburg, Russia (RUS) Th e region of northwestern Russia has a cool maritime Linking SD and DD Decay climate with cool wet summers and long cold winters. Classifi cations through Indicators Major forest tree species include silver birch (Betula of Decay pendula Roth.), Norway spruce (Picea abies (L.) Karst.), To accurately estimate DD and SD population attributes and Eurasian aspen (Populus tremula L.). Th e mean such as biomass or carbon, the FIA program assigns each annual temperature of July is 16.5 °C, while the mean sampled DD and SD individual to a decay class. To temperature of January is 8.5 °C with an annual mean of facilitate linkage of DD and SD decay classes, indicators 3.4 °C. Th e mean annual precipitation is 708 mm. of decay were developed in this study that were assessed for each study dead wood piece. For both DD and SD, Krasnoyarsk, Russia (RUS) there were fi ve decay classes that span a spectrum of Th is region of central Siberia has a moderately cold and decay from fresh mortality to nearly complete decay. continental climate. It belongs to the Angara province of Decay classes are largely qualitative based on the physical the southern taiga and is dominated by Russian larch- appearance and structural integrity of individual dead Scots pine (Larix siberica Ledeb. and Pinus sylvestris L.) wood pieces (USDA 2007, Woundenberg et al. 2010). forests. Th e vegetation period lasts 126 days with a frost- For SD trees the fi ve decay classes were defi ned as: free period of 110 days. Th e mean temperature of July is 16 °C, while the mean temperature of January is -25 °C, Class 1: All limbs and branches are present; the top of the and the overall mean annual temperature is -3.2 °C. Th e crown is still present; all bark remains; sapwood is intact, mean annual precipitation is 361 mm. with minimal decay; heartwood is sound and hard.

Irkutsk, Russia (RUS) Class 2: Th ere are few limbs and no fi ne branches; the Th is region of Eastern Siberia has sites located in the top may be broken; a variable amount of bark remains; area of strictly continental climate with permafrost. sapwood is sloughing with advanced decay; heartwood is Forests were primarily comprised of Scots pine (Pinus sound at base but beginning to decay in the outer part of sylvestris). In mountainous areas Siberian larch (Larix the upper bole. siberica), Siberian white pine (Pinus siberica Ledeb.), and

4 Class 3: Only limb stubs exist; the top is broken; a Based on decades of research into the ecological role variable amount of bark remains; sapwood is sloughing; of detritus in forest ecosystems, we hypothesized that heartwood has advanced decay in upper bole and is the diff erences between SD and DD would be greatest beginning at the base. for intermediate decay classes (i.e., 2 and 3). Given the lack of time in decay class 1, decay reductions are Class 4: Few or no limb stubs remain; the top is broken; likely minimal for both SD and DD. For decay classes a variable amount of bark remains; sapwood is sloughing; 4 and 5, which are based on structural strength, it is heartwood has advanced decay at the base and is likely that SD and DD are similar given that structural sloughing in the upper bole. strength of wood is highly dependent on density. We also hypothesized that the wood density of SD and DD Class 5: No evidence of branches remains; the top is would diverge more in dry climates than in humid ones broken; <20 percent of the bark remains; sapwood is due to moisture limitations in dry climates that inhibit gone; heartwood is sloughing throughout. SD decomposition. Since SD falls and becomes DD, the input of decayed SD-related wood might infl uence For DD trees the fi ve decay classes were defi ned as: the overall density of DD. Since decay classes are largely based on exterior appearances, it is likely that SD in Class 1: Sound, freshly fallen, intact logs with no rot, no dry climates will be denser than DD for a given decay conks present indicating a lack of decay, original color of class because the exterior of SD will continue to degrade wood, no invading roots, fi ne twigs attached with tight (e.g., bark loss, fragmentation of twigs, etc) whereas bark. the interior will remain relatively sound. Conversely, in very wet environments, fragmentation of bark and Class 2: Sound log sapwood partly soft but can’t be cover of mosses and other bryophytes are likely to make pulled apart by hand, original color of wood, no invading the exterior of DD look decayed despite the slow rate of roots, many fi ne twigs are gone and remaining fi ne twigs decomposition of DD in very wet environments. Based have peeling bark. on these hypotheses, a variety of indicators of decay were visually assessed for both SD and DD in this study Class 3: Heartwood is still sound with piece supporting (Table 2). Similarities in indicators of decay among its own weight, sapwood can be pulled apart by hand or SD and DD pieces sampled in this study will suggest is missing, wood color is reddish-brown or original color, appropriate linkage between the SD and DD decay roots may be invading sapwood, only branch stubs are classifi cation systems and subsequent development of SD remaining which cannot be pulled out of log. decay reduction factors based on extensive DD woody decay information. Class 4: Heartwood is rotten with piece unable to support own weight, rotten portions of piece are soft FIELD AND LABORATORY and/or blocky in appearance, a metal pin can be pushed Decay Classes and Indicators of Decay into heartwood, wood color is reddish or light brown, invading roots may be found throughout the log, branch At each study site we sampled SD and DD covering the stubs can be pulled out. full range of decomposition stages, which necessitated the felling of SD trees. To sample indicators of decay, Class 5: Th ere is no remaining structural integrity to selected SD or DD pieces were categorized into one of the piece with a lack of circular shape as rot spreads out fi ve decay classes based on visual characteristics linked across ground, rotten texture is soft and can become to the degree of decomposition that was noted in the powder when dry, wood color is red-brown to dark previous section: the presence of leaves, twigs, branches, brown, invading roots are present throughout, branch percent bark cover on branches and bole, sloughing of stubs and pitch pockets have usually rotten down. wood, collapsing and spreading of DD or SD (indicating the transition from intact elliptical forms to degraded

5 Table 2.—Decay indicators used to determine decay Wood Density and Density Reduction classes for standing and downed dead trees Factors Code Indicator question Foliage Is foliage present? To measure wood density of SD and DD, samples were Twigs Are twigs present? divided into four sections of similar length. Short pieces Branches Are branches present? (e.g., in more advanced decay classes) were divided into Bark_br Is there bark present on the branches? two or three sections. For each section, end diameters Bark_bole Is there bark present on the bole? and length were recorded. From the end of each section, Moss Is there moss present? a disk (a cross section 5 to 10 cm thick) was cut with Lichens Are there lichens present? a chainsaw to determine wood and bark density. To Beetles Is there evidence of beetle presence? determine disk volume, for each cross-section disk we Woodborer Is there evidence of woodborer presence? recorded the outermost diameter, outer wood diameter, Conks Are there conks present? mean longitudinal thickness, circumference covered by Whiterot Is there white rot present? bark, radial thickness of bark, and mean radial depth of Brownrot Is there brown rot present? decay. Bark was separated from wood and the total wet Casehard Does the bole exhibit case hardening? weight of each tissue was determined. Lengths of bark SW_friable Is the sapwood crushable by hand? and wedge-shaped wood subsamples of ~100 g taken HW_friable Is the heartwood crushable by hand? from each disk using a hammer and chisel were weighed. Ants Is there evidence of ants present? Dry weights were recorded after oven drying at 55 °C SW slough Is the sapwood sloughing? to a constant weight (within 0.01 g). For disks that had Collapse Has the bole shape collapsed? not been dried and weighed, dry weights of wood and Scatter Is the bole scattered in the ground? bark were calculated from total wet weights of these Stub_move Do the branch stubs move in the bole? tissues and the ratio of dry-to-wet weight determined from subsamples. Th e density of the bark and wood for each disk was calculated as the ratio of its dry weight shapes), friability or crushability of wood, color of wood, to undecayed volume (g/cm3). Mean bark density and and mobility of branch stubs (Triska and Cromack mean wood density for each tree were calculated by 1980, Graham and Cromack 1982, Sollins 1982, averaging the density of all the cross-section values of Harmon and Sexton 1996). In addition to visual and each tissue for each tree. Recorded values of percent bark structural indicators of decay used by the FIA program, cover for the whole tree and bark circumferential length biological indicators (Table 2) of decomposition, such as for each disk were used to determine the proportion of moss cover, presence of fungal fruiting bodies, and the total tree volume that was bark. Total density per tree presence of insect galleries, were also noted. In addition, was calculated as the summation of each tissue density at most sites we sampled live trees to determine densities multiplied by the proportion of total tree volume of undecayed wood and bark. Dimensions recorded applicable to that tissue. An average density for each for each tree sampled included length, base and top tree species, decay class, and position (i.e., SD or DD) diameters, and current d.b.h. Th e decay classes of both was calculated using total tree density. Trees were also SD and DD used in this study align with the same decay categorized as either hardwood (i.e., angiosperm) or classes used by the FIA program. Please refer to USDA softwood (i.e., gymnosperm) and total density per tree (2010), Woodall et al. (2010), and Woudenberg et was averaged across each category for each decay class al. (2010) for details regarding the SD and DD decay and position. Finally, an average of total density was classifi cation systems used by FIA. calculated for all trees by decay class and position.

6 Th e density reduction factor was calculated as the ratio variance with decreasing mean. Th erefore, assumptions of the average current density to average undecayed of normality and constant variance of residuals appeared density for each species, category, or combination and to be adequately met. Th rough ANOVA testing, three by each decay class and position (i.e., SD versus DD) to results were postulated: 1) No signifi cant diff erence indicate the relative rate that tree density declined for between DD and SD density reduction factors by each scenario. decay classes. Th is would indicate that decay occurred with similar patterns for dead wood in standing or Statistical Analysis lying positions; 2) Th e main eff ects of position and We used analysis of variance (ANOVA) to test how the decay class would both be signifi cant when the density dependent variable density reduction factor varied by the reduction diverged early in the decomposition process independent variables taxa (i.e., species or hardwood/ (i.e., between class 0 and 1). Th is might suggest that softwood), decay class, and position. Th e following one position started with a lower density than the other; model was fi t: 3) A signifi cant interaction of position and decay class. Th is would indicate that one position was decaying to a lesser extent than the other and that density reduction Ykhij = m + k +h+ di + aj+ daij + khij factors were diverging. We tested these hypotheses at where: three separate levels: individual species, softwoods versus hardwoods, and for all species combined. m is the overall mean value of Ykhij, the average density reduction factor of the jth decay class in the To examine why DD and SD might have diff erent ith position in region k and stand h densities and density reduction factors, we compared k is the random eff ect of the region k that was indicators of decay (Table 2) between the two positions. sampled, that adds variability to the value of Y, We computed the relative frequency that indicators were 2 k=region name, k h ~ N(0,  b), k and k’ are present for two positions and the three taxonomic levels independent described above to examine which characteristics might be associated with diff erences in dependent variables. h is the random eff ect of stand h that was sampled, that adds variability to the value of Y, h=stand 2 All statistical tests based on the model outlined above name, h ~ N(0,  b), h and h’ are independent were performed by the MIXED procedure of SAS th di is the eff ect of the i level of the independent (SAS Institute Inc.) using the LSMEANS statement variable position, i=SD or DD which computes least squares means of fi xed eff ects th aj is the eff ect of the j level of the independent and corresponding t and F tests. Statistical tests were variable decay, j= 1, 2, 3, 4 signifi cant if 0.05 > P > 0.01 and highly signifi cant if P < or = 0.01. We emphasize that a minimal amount of da is the interaction of position and decay ij replication was conducted within species, study sites, and khij is the random error term that adds variability decay classes. Replication was limited by the diffi culty in 2 to the value of Y, khij ~ N(0,  ) and khij and fi nding study sites with a full representation of DD and khi’j’ are independent. SD decay classes. Furthermore, the felling of SD trees is extremely hazardous to the sawyer. We have expressed When comparing softwoods and hardwoods, species was the high levels of uncertainty in Appendix C. Th e results added as an additional random variable in the model. In presented in this study should be considered preliminary the case of all species combined, the variables species and to guide future research and promote refi ned SD type (softwood or hardwood) were added as additional population estimation (i.e., use of decay reduction factors random variables. Residuals from the models were fairly in biomass/carbon estimates). symmetric and there was no indication of increasing

7 Development of Standing Dead (SD) Databases A similar uncertainty estimate was made for density We used the fi ndings of the current study to estimate reduction factors using the average density reduction decayed density and the density reduction factors of SD factor and associated uncertainty for DD. for tree species inventoried by the FIA program. For the species that we sampled, we used the observed densities RESULTS and density reduction factors. For the species that we did Decay Indicators not sample, we calculated decayed density and density All Species reduction factors using ratios of the species we sampled and the estimated values for DD from Harmon et al. Th e relative frequency of decay indicators (Table 2) as DD (2008). Specifi cally, for decay classes 1 to 3 we multiplied and SD decomposed was surprisingly similar (Figure 1). the SD to DD ratio for either hardwoods or softwoods In terms of physical indicators (presence of parts, shape, (depending on the species) by the DD density or density friability) prior to reaching decay class 4, DD and SD were reduction factor. For decay class 4, there were few species quite comparable and go through a similar progression sampled. We therefore used the average SD to DD ratio from one decay class to the next. Decay class 4 DD and of all species sampled. For decay class 5, as no species SD diff ered in terms of structural integrity indicators. Th is were sampled given the scarcity of these trees, this study is a logical result as SD that lacks physical integrity tends suggests future research or assuming decay class 4 wood to fall and form DD. Th e nearly entire lack of bark in density results can be extrapolated to decay class 5. decay class 4 SD, as compared to DD, is related to vertical orientation, as even loose bark can remain on the surface To provide an estimate of uncertainty, we used one of of DD. Across all species and study sites, the means of SD the following methods: bark cover (percent of piece covered by bark) were 91.62, 65.55, 38.91, and 21.11 for decay classes 1 through 4, A. Species that had been sampled for SD density. respectively. We used standard error of the observed values of decayed density or the density reduction factor. In terms of biological indicators (presence of moss, lichens, insects, and fungi), the major diff erence between B. Species lacking sampling to determine decayed SD SD and DD appeared to be the abundance of moss. density. Th e uncertainty (U) in the decayed SD DD tended to have higher presence of moss than SD, density for this set of species was estimated by: which would be consistent with the wetter microclimate associated with DD versus SD. Conversely, lichens were slightly more abundant on SD than DD, which is ଶ ଶ ଶ ଶ ୈୣୡୟ୷ୣୢୗୈ ൌ ටሺ‡ୈୈሻሺୈୈሻ൅ሺ ሻሺୖሻ consistent with a drier microclimate associated more with SD than DD. DD also had a greater tendency to support where UDD is the uncertainty in DD (Harmon et al. fungal fruiting bodies, whereas SD had a greater tendency 2008), DenDD is the mean DD density, and UR is the to support wood borers. Th e former may indicate a uncertainty in R, the mean SD to DD density ratio. more favorable decomposition environment; the latter Th is formula accounts for the fact that uncertainty for may be an artifact related to greater moss cover and decayed SD density is a function of two uncertainties. decay state in DD hiding evidence of wood borers. For Our formula assumed no correlation between the further evaluation of these moss/lichen results we suggest uncertainties. Uncertainty was estimated for each decay consulting the abundance of literature regarding lichen class of SD. For decay classes 1 to 3 we used R values substrate preferences. associated with either hardwoods or softwoods and applied them to appropriate taxa. For decay class 4 there We observed little bark cover in both class 2 and 3 SD were few observations, so we used values for all the taxa for some species, making the characteristic less useful as sampled. an indicator than for DD. Two characteristics proved

8 Figure 1.—Percent frequency of abundance for physical indicators of decay class for standing dead and downed dead trees useful in distinguishing these two decay classes when long periods of time, thus the presence/absence of surface SD had little bark cover. First, decay class 3 SD tended biological indicators may not fully indicate internal decay to have more lichens than decay class 2 SD. Th is is dynamics. Barring destructive sampling, standard forest likely related to the older surface having more time to inventory fi eld crew must continue to rely on qualitative accumulate lichens. Second, the wood of decay class SD appearance to determine decay classifi cation. 3 SD tended to take on what might be described as a “ropey” texture. Th is is caused by a weakening of the Hardwoods versus Softwoods outer wood layers causing cracks that look similar to the While the overall relationship between external strands in a rope often associated with white-rot fungus. characteristics and decay classes were similar for Findings by Aakala et al. (2008) indicate that the surface hardwoods and softwoods, there were some notable appearance of standing dead trees can remain static for diff erences (Figure 2). In terms of physical indicators,

9 Figure 2.—Percent frequency of abundance for physical indicators of decay class for hardwood and softwood standing and downed dead trees

hardwoods appeared to lose twigs and branches to a indicated adequate decay class diff erentiation, linkage higher degree than softwoods as decay class increased. between SD and DD decay classifi cation systems, and Conversely, softwoods lost bark on the bole to a higher FIA sampling protocol alignment allowing subsequent degree than hardwoods, although this was likely related development of wood density estimates for SD tree to the presence of birch (Betula sp.), a genus well species in U.S. forests based on relatively extensive DD known for bark retention. For the biological indicators, wood density information. hardwoods were more likely to have white-rot fungus, whereas softwoods were more likely to have brown-rot Wood Density fungus, at least in decay classes 2 and 3. In decay class As expected, SD wood density declined with decay class 1, hardwood SD were more likely to have lichens and (Appendix A). Th e initial, undecayed tree densities for mosses present than softwoods, but this likely stems from the 19 species studied ranged from 0.31 to 0.54 g/cm3 the diff erence of these tree taxa to support epiphytes in (Table 3). Th e overall mean initial density was 0.39 general. g/cm3. Decay class 3, the most advanced decay class common to SD and DD, had densities that ranged from Overall, indicators of decay were not evaluated in 0.16 to 0.34 g/cm3 for DD and 0.21 to 0.46 g/cm3 for this study as an in-depth discourse on SD and DD SD. Decay class 3 density for all species of DD averaged decay dynamics and ecology, rather as a qualitative 0.26 g/cm3 and averaged 0.35 g/cm3 for SD indicating means to assure diff erentiation between decay classes that SD contain less decay than logs (Table 4). Th is is and alignment with FIA sampling protocols. Results

10 Table 3.—Undecayed wood density by tree species code, undecayed density for hardwoods and softwoods was scientifi c name, and location of sample collection (U.S. 3 study site or general Russian location) 0.40 and 0.38 g/cm , respectively. However, decay class 3 hardwood DD and SD had mean densities of 0.23 and Undecayed 3 Density 0.24 g/cm , whereas softwood DD and SD had mean Code Scientifi c name Location (g/cm3) densities of 0.27 and 0.38 g/cm3, respectively. PILU Picea x lutzii AK 0.36 PICO Pinus contorta FEF 0.40 Density Reduction Factors PIBA2 Pinus banksiana MEF 0.42 Individual species POTR Populus tremuloides MEF 0.39 LITU Liriodendron tulipifera NCB 0.41 Density reduction factors for decay class 3 ranged from PINE Pinus species NCB 0.49 0.34 to 0.80 and from 0.44 to 1.11 for DD and SD, TSHE Tsuga heterophylla CHE 0.44 respectively (Appendix B). Statistical analysis showed fi ve ABSI Abies siberica RUS 0.31 species with highly signifi cant interactions and two with BECO Betula costata RUS 0.54 signifi cant interactions between position and decay class BEPE Betula pendula RUS 0.48 (P-values: Larix sibirica 0.00; Pinus contorta 0.00; Picea LADA Larix daurica RUS 0.46 lutzii 0.00; Pinus species 0.01; Pinus sibirica 0.01; Pinus LASI Larix sibirica RUS 0.49 banksiana 0.05; Pinus koraiensis 0.05). Of these seven PIAB Picea abies RUS 0.37 species, four were in the genus Pinus. Th is interaction PIAJ Picea ajanensis RUS 0.36 indicated that SD for these species retained a higher PIOB Picea obovata RUS 0.38 density than DD and that these diff erences increased PIKO Pinus koraiensis RUS 0.35 as decay proceeded. Aakala (2010) found similar PISI2 Pinus sibirica RUS 0.34 results in Picea abies forests of northern Europe, with PISY Pinus sylvestris RUS 0.38 the wood density of SD signifi cantly higher than DD. POTR2 Populus tremula RUS 0.35 Furthermore, in our study three species (two hardwood and one softwood) had highly signifi cant decay class and position main eff ects, but no signifi cant interaction Table 4.—Average density (g/cm3) for standing dead (SD) to downed dead (DD) trees and associated ratio (SD/DD between the two factors. Th is result may indicate that average density) for all species and sites combined by early in the decay process, DD and SD diverge, with decay class DD undergoing an initial rapid period of decomposition Decay SD DD SD/DD and SD subject to a lag. Four species (three hardwoods class (g/cm3)n(g/cm3)nRatio and one softwood) showed only decay class as a highly 1 0.40 (0.00) 175 0.40 (0.00) 185 1.00 signifi cant factor in density reduction (P-values: Betula 2 0.38 (0.00) 189 0.33 (0.00) 292 1.15 3 0.35 (0.01) 78 0.26 (0.00) 391 1.35 pendula 0.00; Tsuga heterophylla 0.00; Populus tremula 4 0.25 (0.03) 14 0.15 (0.00) 246 1.67 0.00; Populus tremuloides 0.00). Th is would indicate 5 0.11 (0.00) 174 that decay progresses similarly in SD and DD for these Note: Values are means with standard errors in parentheses. “n” is species. Two species showed only position as a highly the number of SD or DD. signifi cant factor in density reduction, but this was likely an artifact related to the few decay classes sampled for consistent with the notion that SD must retain some these species. sound wood to remain upright. Aakala’s (2010) study in Picea abies forests in northern Europe found a range of Hardwoods versus softwoods SD decayed density ranging from 0.14 to 0.47 g/cm3. Compared to hardwoods, softwoods, on average, are more likely to have a greater divergence in SD and DD Hardwoods tended to have slightly higher initial decay reduction factors as decay progresses (Table 6). density than softwoods, but the opposite was true after For example, the mean density reduction factors for extensive decay (Table 5). For example, the mean initial, decay class 3 hardwood DD and SD were 0.51 and 0.54,

11 Table 5.—Average density (g/cm3) for standing dead (SD) and downed dead (DD) trees and associated ratio (SD/DD average density) by hardwood/softwood and decay class Type Decay SD DD Ratio SD/ class (g/cm3)n (g/cm3)nDD Hardwood 1 0.46 (0.01) 37 0.43 (0.01) 51 1.07 2 0.36 (0.01) 31 0.33 (0.01) 58 1.09 3 0.24 (0.02) 14 0.23 (0.01) 60 1.04 4 0.20 (0.03) 2 0.13 (0.01) 55 1.54 5 0.11 (0.01) 31 Softwood 1 0.38 (0.00) 138 0.38 (0.00) 134 1.00 2 0.39 (0.00) 158 0.34 (0.00) 234 1.15 3 0.38 (0.01) 64 0.27 (0.00) 331 1.41 4 0.26 (0.03) 12 0.15 (0.00) 191 1.73 5 0.11 (0.00) 143 Note: Values are means with standard errors in parentheses. “n” is the number of SD or DD.

Table 6.—Average density reduction factor (decayed density/undecayed density) for standing dead (SD) and downed dead (DD) trees and associated ratio (SD/DD density reduction factor) by hardwood/softwood and decay class Type Decay SD n DD n Ratio SD/ class DD Hardwood 1 0.99 (0.01) 37 0.95 (0.01) 51 1.04 2 0.80 (0.02) 31 0.74 (0.02) 58 1.08 3 0.54 (0.04) 14 0.51 (0.03) 60 1.06 4 0.43 (0.05) 2 0.29 (0.02) 55 1.48 5 0.22 (0.02) 31 Softwood 1 0.97 (0.01) 138 0.93 (0.01) 134 1.04 2 1.00 (0.01) 158 0.87 (0.01) 234 1.15 3 0.92 (0.03) 64 0.70 (0.01) 331 1.33 4 0.55 (0.06) 12 0.40 (0.01) 191 1.38 5 0.29 (0.01) 143 Note: Values are means with standard errors in parentheses. “n” is the number of SD or DD.

respectively. In contrast, the mean density reduction was a highly signifi cant interaction between position and factors for decay class 3 softwood DD and SD were 0.70 decay class in the reduction factor, which may be due to and 0.92, respectively. For softwood species collectively, softwood species being over represented in the data set. there was a highly signifi cant interaction between position and decay class eff ects. For hardwood species, Standing Dead Databases decay class was a highly signifi cant factor in density Of 260 species considered by the FIA inventory, eight reduction. Th is indicates that DD density factors could (3 percent) have had some SD decay classes sampled. be substituted for SD ones in hardwoods, but not for For sampled species, the uncertainty of SD density was softwoods. approximately 3 to 14 percent of the mean estimate. In contrast for species in which SD density had to be All species combined estimated from the DD database produced by Harmon Th e density reduction factors for all decay class 3 DD et al. (2008), the uncertainty is approximately 20 to 100 and SD was 0.67 and 0.86, respectively (Table 7). Th ere

12 Table 7.—Average density reduction factor for standing 2009). Th is might lead to consistent diff erences in the dead (SD) and downed dead (DD) trees and associated ratio (SD/DD density reduction factor) for all sites/ density of SD versus DD. In very humid environments, species combined by decay class DD and SD both may retain moisture and hence the Decay Ratio SD/ density of SD and DD might remain similar. Th is may class SD n DD n DD explain our results from Cascade Head, the wettest site 1 0.97 (0.01) 175 0.94 (0.01) 185 1.03 examined. In very dry environments SD decay may be 2 0.97 (0.01) 189 0.84 (0.01) 292 1.15 limited relative to that of DD and that could lead to an 3 0.86 (0.03) 78 0.67 (0.01) 391 1.28 increasing diff erence as decay class increases. Specifi c to 4 0.53 (0.06) 14 0.38 (0.01) 246 1.39 sites in this study, but possibly not pertinent to all forest 5 0.28 (0.01) 174 conditions across the United States, softwoods tended to Note: Values are means with standard errors in parentheses. “n” is the number of SD or DD. grow in drier climates and microsites than hardwoods, which may explain why many softwoods exhibited an increasing diff erence in density as decay classes increased. percent of the mean estimate, the latter for decay class In environments with intermediate moisture, SD may 4 SD. Th is indicates that actual sampling of SD density be slightly drier than DD. Th is could lead to a relatively could reduce the uncertainties with this material at least short lag in decomposition that would result in a an order of magnitude. constant diff erence between decay classes in terms of Implications of Density Reduction on density. While our observations are consistent with these Population Estimates hypotheses, other factors could be causing the patterns observed and more SD:DD comparisons in a wider Th e implications of assumptions of SD density can be range of environments would be helpful in determining potentially profound in terms of biomass estimates. the mechanisms explaining these phenomena. Using FIA’s forest inventory of the state of Minnesota Diff erences in the density of SD versus DD may also be (2005-2009), we explored the impact of three plausible related to the physical limits of strength of woody tissues. scenarios for estimating total SD biomass (i.e., SD wood Th at is, SD have to be fully self supporting, and once density assumptions): 1) SD wood density equals live their strength decreases past a threshold they will collapse tree wood density; 2) SD and DD wood density are to form DD. In contrast, DD with minimal structural equal; and 3) SD wood density based on this study’s strength can collapse, but remain as DD. Th is may lead newly created SD density reduction factors. To convert to an apparent abundance of higher wood density SD that volumetric estimate to one of biomass, the three SD trees as less sound SD fall to become DD. However, wood density assumptions were evaluated. Assuming SD do not necessarily have to decompose throughout that SD wood density was equal to live trees gives a total to fall. For example, SD in dry environments fall after SD biomass estimate of 32.98 million Mg. By assuming root systems decompose, adding relatively sound wood that SD and DD wood density are equal, the estimated to the DD pool. When this occurs the density of DD biomass was 24.38 million Mg, or 26 percent less. Using might increase due to this input of relatively sound SD the new estimates of SD wood density provided an wood. Th erefore, the diff erence in wood density between estimated biomass of 27.52 million Mg or 16.5 percent SD and DD are likely to be controlled by a number of less than the assumption of undecayed density. interacting factors. Care should be taken not to infer rates of decay with assigned decay classes. Diff erences in DISCUSSION wood density were quantitively determined between SD SD inhabit a very diff erent environment than DD, with and DD qualitative decay classes in this study. Rates of the former usually being drier. Given that the response decay are only inferred as initial hypotheses for future of decomposition to moisture shows an optimum research. range (i.e., woody tissues can be too dry or too wet to decompose readily), the position supporting the fastest Th ere is an emerging abundance of research discussing decomposition can change with macroclimate (Harmon the dynamics of SD and DD decay and subsequent

13 eff ects on biomass/residence times (Aakala et al. 2008, develop a SD decay class system based on a wider range Aakala 2010, Kruys et al. 2002, Vanderwel et al. 2006, of species better aligned with the DD system. Zielonka 2006,). Moving beyond initial characterizations of SD and DD decay dynamics on intensively monitored While it is important to eventually determine density and study sites towards estimates of SD wood density by density reduction factors for SD and DD of the major decay class for all forest tree species in the United States species encountered in inventories, in the short-term one is a tremendous change in study scope. However, the is forced to make assumptions when species or positions immediate need to develop and document SD decay have not been sampled. Whether considering individual reduction factors for most forest tree species in the species, hardwoods and softwoods, or all samples United States to meet mandated carbon stock reports combined, the ranges of density reduction factors for SD requires a timely assimilation of the best available science. were consistently higher compared to DD. Th is indicates Given the lack of information on SD decayed wood generally less reduction in density as decay progresses in density for hundreds of tree species in the United States, SD than DD. Softwood species tended to exhibit this several approaches were considered. First, SD tree density pattern more strongly than hardwood species, sometimes can be assumed to be similar to undecayed tree density. even increasing in density. Th ere were exceptions to this For some environments in which decomposition is very pattern in the statistical analysis (insignifi cant eff ect of slow, this might be acceptable. However, in our analysis position), but inspection of the charted data indicated we did not fi nd a single species in which density did not that these cases had samples for only decay classes 1 and decline with decay class. Second, SD tree density could 2, were hardwood species, or were located in a warm and be assumed equal to that of DD tree density. Th is would wet climate. Th e assumption with the strongest support be appropriate for 6 of the 19 species we examined. at this time is that softwoods tend to have a greater Th e most common situation in our study (13 of 19 divergence in SD and DD density reduction factors than species) was for either position or a position X decay class hardwoods. However, additional sampling of species, interaction to be signifi cant. hardwoods in particular, is required to confi rm if this is actually the case. To create the needed SD decay reduction factors for all tree species in the United States, decay classes in this Irrespective of the fi ndings of this study, SD wood study were aligned with those used by FIA using this density at some point in its decay process will become study’s indicators of decay. FIA’s SD decay classes 1 substantially less than that of living trees. Th e selection and 2 were quite similar to the classes 1 and 2 assessed of density reduction factors can have a profound eff ect in this study. Given the more highly decayed nature of on resulting population estimates (e.g., SD carbon in SD decay classes 3 and 4, FIA’s SD decay classes 3 and a state). Th is study found substantial variation in SD 4 seem to span the characteristics of decay class 3 SD biomass population estimates for Minnesota when using trees with some overlap into decay class 4. Th e FIA SD undecayed density versus reduced wood density values. decay class 5 would seem to be similar to this study’s SD Th e diff erences in estimates depend strongly on the decay class 4. Based on this fi nding, it is suggested that dominance of hardwoods versus softwoods. When only the decay reductions for SD decay class 4 be used as an softwood species were considered, the diff erence between approximation for SD decay class 5 until such time that the assumption of live tree wood density and the SD empirical information refi nes SD decay class 5 attributes densities that we estimated was 2.5 percent. In contrast, or a model is suggested to extrapolate decay class 4 when only hardwood species were considered the diff erence attributes to decay class 5 pieces. Furthermore, given the between these two methods was 22 percent. Th erefore, the potential safety hazard of felling decay class 5 SD trees, potential biases contained in current SD estimates are likely this assumption may be the most practical. In general, to vary substantially. Th is analysis is considered preliminary these are gross approximations and, given the fact that for illustrative purposes as structural reductions for decay the FIA system is based on Douglas-fi r with unspecifi ed were not incorporated into the estimation procedures. modifi cations for other species, it would be logical to

14 CONCLUSIONS LITERATURE CITED To provide accurate, unbiased estimates of the biomass Aakala, T. 2010. Coarse woody debris in late- and carbon stocks of SD trees, it is necessary to have successional Picea abies forests in northern Europe: species- and decay class-specifi c estimates of either Variability in quantities and models of decay class density or density reduction factors. Empirically dynamics. Forest Ecology and Management. 260: sampling the wood density of SD by species and decay 770-779. class across the forests of the United States is currently restricted by the inherent costs and felling hazards Aakala, T.; Kuuluvainen, T.; Gauthier, S.; DeGrandpre, involved. As an approach to estimate preliminary SD L. 2008. Standing dead trees and their decay-class wood density reduction factors, this study established dynamics in the northeastern boreal old-growth relationships between SD and DD such that extensive forests of Quebec. Forest Ecology and Management. DD woody density studies could be used to suggest 255: 410-420. SD wood density attributes. Th is study suggests a new set of SD decay specifi c density and density reduction Adams, M.B.; Loughry, L.; Plaugher, L., comps. 2004. factors for many tree species in the United States that Experimental Forests and Ranges of the USDA may be used to refi ne FIA’s SD population estimation Forest Service. Gen. Tech. Rep. NE-321. Newtown procedures. Given the size of the uncertainty involved Square, PA: U.S. Department of Agriculture, Forest with the SD wood decay attributes proposed in this Service, Northeastern Research Station. 178 p. study, additional work to develop a multispecies SD decay class system and to better characterize diff erences Dunn, J. 1977. Soil survey of Orange County, NC. between SD and DD trees is recommended. Raleigh, NC: U.S. Department of Agriculture, Soil Conservation Service. 93 p. ACKNOWLEDGMENTS Th is work was funded by a joint venture agreement Fasth, B.; Harmon, M.E.; Sexton, J.; White, P.S. 2011. between the U.S. Forest Service Forest Inventory and In press. Decomposition of fi ne woody debris in a Analysis and Oregon State University (09-JV-11242305- deciduous forest in North Carolina. Journal of the 025). Torrey Botanical Society.

Special thanks are extended to Shawn Fraver, Ph.D., Franklin, J.F.; Shugart, H.H.; Harmon, M.E. 1987. Tree and Brad Oberle, Ph.D., who provided abundant death as an ecological process. BioScience. 37(8): constructive criticism that improved the quality of this 550-556. documentation. Graham, R.L.; Cromack, K. 1982. Mass, nutrient Th e cooperation of numerous experimental forest staff at content, and decay rate of dead boles in rain forests locations in the United States and Russia were critical to of Olympic National Park. Canadian Journal of this study’s success. In particular, Randy Kolka, Ph.D. Forest Research. 12: 511-521. and his staff at the Marcell Experimental Forest were especially helpful with facilitating the fi eld work. We Greene, S.; Acker, S. 1999. Growth and Yield would also like to thank Peter White, Ph.D. of the North Permanent Plots at the Cascade Head Experimental Carolina Botanical Garden for the use of facilities and his Forest. Available at http://andrewsforest.oregonstate. many students whose eff orts made this project possible. edu/pubs/webdocs/reports/permplot/psp3. cfm?topnav=55 (Accessed June 13, 2011).

15 Harmon, M.E. 2009. Woody detritus mass and its Norway spruce (Picea abies) trees in managed contribution to carbon dynamics of old-growth Swedish boreal forests. Canadian Journal of Forest forests: Th e temporal context. In: Wirth, C.; Research. 29: 178-186. Gleixner, G.; Heimann, M. eds. Old-growth forests: Function, fate and value. New York. Springer-Verlag Kruys, N.; Jonsson, B.G.; Stahl, G. 2002. A stage-based Ecological Studies. 207: 159-189. matrix model for decay-class dynamics of woody debris. Ecological Applications. 12: 773-781. Harmon, M.E.; Sexton, J. 1996. Guidelines for measurements of woody detritus in forest ecosystems. Lee, P. 1998. Dynamics of snags in aspen-dominated US LTER Publication No. 20. Seattle, WA: U.S. LTER midboreal forests. Forest Ecology and Management. Network Offi ce, University of Washington, College of 105: 263-272. Forest Resources, Seattle, WA. 73 p. Miles, P.D.; Smith, W.B. 2009. Specifi c gravity Harmon, M.E.; Fasth, B.; Yatskov, M.; Sexton, J. and other properties of wood and bark for 156 2005. Th e fate of beetle-killed spruce on the Kenai tree species found in North America. Res. Note. Peninsula: A preliminary report. Gen. Tech. Rep. NRS-38. Newtown Square, PA: U.S. Department R10-TP-134. Anchorage, AK: U.S. Department of of Agriculture, Forest Service, Northern Research Agriculture, Forest Service State and Private Forestry. Station. 35 p. 23 p. Nilsson, S.G.; Hedin, J.; Niklasson, M. 2001. Harmon, M.E.; Woodall, C.W.; Fasth, B.; Sexton, J. Biodiversity and its assessment in boreal and 2008. Woody detritus density and density reduction nemoral forests. Scandinavian Journal of Forest factors for tree species in the United States: a Research. 3(suppl): 10-26. synthesis. Gen. Tech. Rep. NRS-29. Newtown Square, PA: U.S. Department of Agriculture, Forest Smithwick, E.A.H.; Harmon, M.E.; Remillard, S.M.; Service, Northern Research Station. 84 p. Acker, S.A.; Franklin, J.F. 2002. Potential upper bounds of carbon stores in forests of the Pacifi c Harmon, M.E.; Franklin, J.F.; Swanson, F.J.; Sollins, P.; Northwest. Ecological Applications. 12: 1303-1317. Lattin, J.D.; Anderson, N.H.; Gregory, S.V.; Cline, S.P.; Aumen, N.G.; Sedell, J.R.; Lienkaemper, G.W.; Sollins, P. 1982. Input and decay of coarse woody Cromack, K. Jr.; Cummins, K.W. 1986. Th e ecology debris in coniferous stands in western Oregon and of coarse woody debris in temperate ecosystems. Washington. Canadian Journal of Forest Research. Recent Advances in Ecological Research. 15: 133-302. 12: 18-28.

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Jonsell, M.; Weslien, J.; Ehnstrom, B. 1998. Substrate U.S. Department of Agriculture, Forest Service. 2007. requirements of red-listed saproxylic invertebrates Forest Inventory and Analysis national core fi eld in Sweden. Biodiversity Conservation. 7: 749-764. guide, volume I: Field data collection procedures for phase 2 plots, version 4.0. Washington, DC: Kruys, N.; Fries, C.; Jonsson, B.G.; Lamas, T.; Stahl, U.S. Department of Agriculture Forest Service, G. 1999. Wood inhabiting cryptograms on dead Forest Inventory and Analysis, Washington, D.C.:

16 Available at: http://www.fi a.fs.fed.us/library/ (Accessed Inventory and Analysis Database Version 4.0: December 2010). Description and users manual for Phase 3. Gen. Tech. Rep. NRS-61. Newtown Square, PA: U.S. Vanderwel, M.; Malcolm, J.R.; Smith, S.M. 2006. An Department of Agriculture, Forest Service, Northern integrated model for snag and downed woody Research Station. 180 p. debris decay class transitions. Forest Ecology and Management. 234: 48-59. Woudenberg, S.W.; Conkling, B.L.; O’Connell, B.M.; LaPoint, E.B.; Turner, J.A.; Waddell, K.L. 2010. Th e Woodall, C.W., Monleon, V.J. 2008. Sampling Forest Inventory and Analysis database: Database protocol, estimation, and analysis procedures for description and users manual version 4.0 for Phase the down woody materials indicator of the FIA 2. Gen. Tech. Rep. RMRS-GTR- 245. Fort Collins, program. Gen. Tech. Rep. NRS-22. Newtown CO: U.S. Department of Agriculture, Forest Service, Square, PA: U.S. Department of Agriculture, Forest Rocky Mountain Research Station. 339 p. Service, Northern Research Station. 68 p. Yatskov, M.A. 2000. A chronosequence of wood Woodall, C.W.; Rondeux, J.; Verkerk, P.J.; Ståhl, G. decomposition in the boreal forests of Russia. 2009. Estimating dead wood during national forest Corvallis, OR: Oregon State University. 168 p. M.S. inventories: A review of inventory methodologies thesis. and suggestions for harmonization. Environmental Management. 44: 624-631. Zielonka, T. 2006. Quantity and decay stages of coarse woody debris in old-growth subalpine forests of Woodall, C.W.; Conkling, B.L.; Amacher, M.C.; western Carpathians, Poland. Canadian Journal of Coulston, J.W.; Jovan, S.; Perry, C.H.; Schulz, Forest Research. 36: 2614-2622. B.; Smith, G.C.; Will-Wolf, S. 2010. Th e Forest

17 Appendix A Average density (g/cm3) for standing dead (SD) and downed dead (DD) trees and their associated ratio (SD/DD average density) for all study species Species Decay SD DD Ratio code class (g/cm3)n(g/cm3)nSD/DD ABSI 1 0.30 (0.02) 7 ABSI 2 0.33 (0.02) 4 0.30 (0.02) 4 1.10 ABSI 3 0.32 () 1 0.25 (0.03) 7 1.28 ABSI 4 0.20 (0.02) 10 ABSI 5 0.17 (0.02) 7 BECO 1 0.54 (0.01) 5 0.50 (0.01) 8 1.08 BECO 2 0.33 (0.04) 3 BECO 3 0.24 (0.00) 2 0.18 (0.01) 8 1.33 BECO 4 0.12 (0.00) 2 BECO 5 0.08 (0.00) 2 BEPE 1 0.48 (0.01) 21 0.47 (0.01) 25 1.02 BEPE 2 0.39 (0.01) 19 0.37 (0.01) 28 1.05 BEPE 3 0.27 (0.03) 6 0.25 (0.02) 29 1.08 BEPE 4 0.20 (0.03) 2 0.15 (0.01) 41 1.33 BEPE 5 0.11 (0.01) 29 LADA 1 0.44 (0.01) 2 0.46 (0.01) 13 0.96 LADA 2 0.46 (0.03) 8 0.39 (0.02) 13 1.18 LADA 3 0.33 (0.02) 14 LADA 4 0.19 (0.01) 9 LADA 5 0.11 (0.02) 4 LASI 1 0.45 (0.01) 12 0.46 (0.01) 16 0.98 LASI 2 0.46 (0.02) 11 0.40 (0.01) 13 1.15 LASI 3 0.46 (0.01) 12 0.34 (0.02) 16 1.35 LASI 4 0.15 (0.01) 16 LASI 5 0.11 (0.01) 16 LITU 0 0.40 (0.00) 2 LITU 1 0.37 (0.01) 6 0.31 (0.05) 3 1.19 LITU 2 0.35 (0.01) 5 0.28 (0.02) 9 1.25 LITU 3 0.20 (0.03) 5 LITU 4 0.11 (0.02) 3 PIAB 1 0.37 (0.03) 4 0.33 (0.01) 5 1.12 PIAB 2 0.37 (0.02) 5 0.27 (0.01) 6 1.37 PIAB 3 0.21 (0.02) 7 PIAB 4 0.13 (0.05) 3 PIAJ 1 0.38 (0.01) 8 0.34 (0.01) 11 1.12 PIAJ 2 0.39 (0.01) 3 0.32 (0.02) 11 1.22 PIAJ 3 0.36 () 1 0.24 (0.02) 15 1.50 PIAJ 4 0.15 (0.02) 10 PIAJ 5 0.12 (0.01) 6 PIBA2 1 0.39 (0.02) 7 0.39 (0.01) 5 1.00 PIBA2 2 0.38 (0.01) 5 0.36 (0.01) 7 1.06 PIBA2 3 0.32 (0.04) 6 0.22 (0.02) 6 1.45 PIBA2 4 0.12 (0.01) 7 PICO 1 0.39 (0.00) 27 0.37 (0.01) 18 1.05 PICO 2 0.41 (0.01) 8 0.36 (0.01) 15 1.14 continued

18 Appendix A.—continued Species Decay SD DD Ratio code class (g/cm3)n(g/cm3)nSD/DD PICO 3 0.40 (0.00) 7 0.32 (0.02) 15 1.25 PICO 4 0.17 (0.01) 7 PICO 5 0.16 (0.01) 9 PIKO 1 0.36 (0.00) 4 0.35 (0.01) 8 1.03 PIKO 2 0.34 (0.02) 5 0.32 (0.01) 10 1.06 PIKO 3 0.33 (0.02) 3 0.24 (0.01) 12 1.38 PIKO 4 0.16 (0.01) 4 PIKO 5 0.13 (0.03) 2 PILU 0 0.38 (0.01) 11 PILU 1 0.38 (0.01) 21 0.36 (0.01) 7 1.06 PILU 2 0.38 (0.01) 69 0.34 (0.01) 92 1.12 PILU 3 0.40 (0.02) 17 0.26 (0.01) 153 1.54 PILU 4 0.13 (0.01) 58 PILU 5 0.08 (0.00) 52 PINE 1 0.37 (0.02) 5 0.37 (0.03) 5 1.00 PINE 2 0.36 (0.02) 4 0.27 (0.02) 9 1.33 PINE 3 0.35 (0.05) 5 0.19 (0.02) 8 1.84 PINE 4 0.35 (0.03) 6 0.19 (0.03) 7 1.84 PINE 5 0.22 () 1 PIOB 1 0.36 (0.02) 4 0.37 (0.01) 10 0.97 PIOB 2 0.39 (0.01) 4 0.33 (0.02) 10 1.18 PIOB 3 0.25 (0.01) 14 PIOB 4 0.13 (0.01) 13 PIOB 5 0.10 (0.01) 8 PISI2 1 0.33 (0.01) 8 0.33 (0.01) 8 1.00 PISI2 2 0.34 (0.00) 7 0.30 (0.01) 8 1.13 PISI2 3 0.30 (0.00) 2 0.23 (0.02) 8 1.30 PISI2 4 0.14 (0.01) 8 PISI2 5 0.10 (0.01) 8 PISY 1 0.37 (0.01) 23 0.36 (0.01) 20 1.03 PISY 2 0.36 (0.01) 19 0.32 (0.01) 31 1.13 PISY 3 0.31 (0.03) 4 0.27 (0.01) 47 1.15 PISY 4 0.18 (0.01) 32 PISY 5 0.12 (0.01) 29 POTR 1 0.37 (0.02) 5 0.36 (0.01) 7 1.03 POTR 2 0.29 (0.02) 6 0.28 (0.02) 6 1.04 POTR 3 0.21 (0.03) 5 0.16 (0.01) 7 1.31 POTR 4 0.07 (0.01) 6 POTR2 1 0.35 (0.01) 8 POTR2 2 0.25 () 1 0.29 (0.02) 12 0.86 POTR2 3 0.24 () 1 0.25 (0.02) 11 0.96 POTR2 4 0.14 (0.01) 3 TSHE 1 0.39 (0.01) 6 0.42 (0.01) 8 0.93 TSHE 2 0.36 (0.02) 6 0.35 (0.03) 5 1.03 TSHE 3 0.29 (0.02) 6 0.28 (0.02) 9 1.04 TSHE 4 0.17 (0.02) 6 0.17 (0.03) 7 1.00 TSHE 5 0.12 () 1 Note: Values are means with standard errors in parentheses. “n” is the number of SD or DD.

19 Appendix B Average density reduction factor for standing dead (SD) and downed dead (DD) trees and their associated ratio (SD/DD decay reduction factor) for all study species Species Ratio code Decay class SD n DD n SD/DD ABSI 1 0.95 (0.06) 7 ABSI 2 1.06 (0.06) 4 0.97 (0.05) 4 1.09 ABSI 3 1.03 () 1 0.79 (0.08) 7 1.30 ABSI 4 0.66 (0.07) 10 ABSI 5 0.55 (0.08) 7 BECO 1 1.00 (0.01) 5 0.93 (0.01) 8 1.08 BECO 2 0.62 (0.07) 3 BECO 3 0.44 (0.01) 2 0.34 (0.02) 8 1.29 BECO 4 0.22 (0.01) 2 BECO 5 0.16 (0.00) 2 BEPE 1 1.01 (0.01) 21 0.98 (0.02) 25 1.03 BEPE 2 0.81 (0.03) 19 0.77 (0.02) 28 1.05 BEPE 3 0.56 (0.07) 6 0.53 (0.04) 29 1.06 BEPE 4 0.43 (0.05) 2 0.30 (0.03) 41 1.43 BEPE 5 0.23 (0.02) 29 LADA 1 0.88 (0.01) 2 0.90 (0.02) 13 0.98 LADA 2 0.92 (0.05) 8 0.77 (0.03) 13 1.19 LADA 3 0.66 (0.04) 14 LADA 4 0.37 (0.03) 9 LADA 5 0.22 (0.05) 4 LASI 1 0.92 (0.02) 12 0.94 (0.03) 16 0.98 LASI 2 0.95 (0.03) 11 0.82 (0.02) 13 1.16 LASI 3 0.95 (0.02) 12 0.70 (0.04) 16 1.36 LASI 4 0.30 (0.02) 16 LASI 5 0.22 (0.02) 16 LITU 0 0.97 (0.00) 2 LITU 1 0.91 (0.03) 6 0.76 (0.12) 3 1.20 LITU 2 0.85 (0.02) 5 0.68 (0.04) 9 1.25 LITU 3 0.48 (0.07) 5 LITU 4 0.28 (0.05) 3 PIAB 1 1.01 (0.09) 4 0.88 (0.03) 5 1.15 PIAB 2 1.01 (0.04) 5 0.73 (0.03) 6 1.38 PIAB 3 0.57 (0.05) 7 PIAB 4 0.36 (0.12) 3 PIAJ 1 1.07 (0.02) 8 0.97 (0.03) 11 1.10 PIAJ 2 1.11 (0.02) 3 0.92 (0.05) 11 1.21 PIAJ 3 1.03 () 1 0.68 (0.06) 15 1.51 PIAJ 4 0.43 (0.05) 10 PIAJ 5 0.34 (0.02) 6 PIBA2 1 0.95 (0.04) 7 0.95 (0.03) 5 1.00 PIBA2 2 0.92 (0.03) 5 0.87 (0.03) 7 1.06 PIBA2 3 0.78 (0.09) 6 0.53 (0.04) 6 1.47 PIBA2 4 0.29 (0.03) 7 PICO 1 0.98 (0.01) 27 0.95 (0.02) 18 1.03 PICO 2 1.04 (0.03) 8 0.92 (0.02) 15 1.13 continued

20 Appendix B.—continued Species Ratio code Decay class SD n DD n SD/DD PICO 3 1.02 (0.01) 7 0.80 (0.05) 15 1.28 PICO 4 0.43 (0.03) 7 PICO 5 0.42 (0.02) 9 PIKO 1 0.92 (0.01) 4 0.90 (0.02) 8 1.02 PIKO 2 0.88 (0.05) 5 0.81 (0.03) 10 1.09 PIKO 3 0.86 (0.05) 3 0.62 (0.03) 12 1.39 PIKO 4 0.41 (0.03) 4 PIKO 5 0.34 (0.07) 2 PILU 0 1.06 (0.03) 11 PILU 1 1.06 (0.02) 21 1.01 (0.04) 7 1.05 PILU 2 1.06 (0.02) 69 0.93 (0.01) 92 1.14 PILU 3 1.11 (0.04) 17 0.73 (0.02) 153 1.52 PILU 4 0.37 (0.01) 58 PILU 5 0.23 (0.01) 52 PINE 1 0.76 (0.05) 5 0.75 (0.06) 5 1.01 PINE 2 0.74 (0.03) 4 0.55 (0.04) 9 1.35 PINE 3 0.72 (0.09) 5 0.40 (0.03) 8 1.80 PINE 4 0.72 (0.07) 6 0.39 (0.06) 7 1.85 PINE 5 0.46 () 1 PIOB 1 0.97 (0.04) 4 0.98 (0.03) 10 0.99 PIOB 2 1.03 (0.03) 4 0.87 (0.05) 10 1.18 PIOB 3 0.66 (0.03) 14 PIOB 4 0.35 (0.03) 13 PIOB 5 0.26 (0.02) 8 PISI2 1 0.86 (0.02) 8 0.85 (0.02) 8 1.01 PISI2 2 0.89 (0.01) 7 0.77 (0.03) 8 1.16 PISI2 3 0.78 (0.01) 2 0.58 (0.04) 8 1.34 PISI2 4 0.36 (0.03) 8 PISI2 5 0.27 (0.02) 8 PISY 1 0.99 (0.02) 23 0.96 (0.02) 20 1.03 PISY 2 0.98 (0.02) 19 0.87 (0.02) 31 1.13 PISY 3 0.83 (0.08) 4 0.71 (0.02) 47 1.17 PISY 4 0.48 (0.04) 32 PISY 5 0.33 (0.02) 29 POTR 1 0.97 (0.04) 5 0.93 (0.03) 7 1.04 POTR 2 0.75 (0.05) 6 0.72 (0.04) 6 1.04 POTR 3 0.54 (0.07) 5 0.41 (0.03) 7 1.32 POTR 4 0.18 (0.01) 6 POTR2 1 0.95 (0.02) 8 POTR2 2 0.66 () 1 0.78 (0.05) 12 0.85 POTR2 3 0.63 () 1 0.66 (0.07) 11 0.95 POTR2 4 0.36 (0.03) 3 TSHE 1 0.90 (0.03) 6 0.96 (0.02) 8 0.94 TSHE 2 0.83 (0.04) 6 0.80 (0.06) 5 1.04 TSHE 3 0.66 (0.05) 6 0.64 (0.05) 9 1.03 TSHE 4 0.38 (0.05) 6 0.40 (0.06) 7 0.95 TSHE 5 0.27 () 1 Note: Values are means with standard errors in parentheses. “n” is the number of SD or DD.

21 continued ABAMABBA 0.400ABBR 0.356 0.340ABCO 0.360 0.020 0.356ABFR 0.370 0.367 0.024 BABGR 0.337 0.340 0.024ABLA B 0.350 0.382 0.012 0.367 BABMA 0.338 0.014 0.414 0.310 0.024 BABMA2 0.360 0.010 0.354 0.014 0.367 B B 0.360 0.473 0.319 0.029 0.014ABPR B B 0.367 0.301 0.012 0.013 0.354ABIE B B 0.024 0.370 0.338 0.060 0.412 0.029ACSP2 B B 0.364 0.013 0.340 0.301 0.058 0.331 BACBA 0.600 B B 0.010 0.435 0.172 0.367 0.060 0.025ACGL 0.565 B B 0.354 0.540 0.239 0.023 0.057 0.301 0.024 BACGR 0.126 B B 0.029 0.440 0.568 0.320 0.336 0.286 0.060ACMA B B B 0.440 0.568 0.309 0.048 B 0.057 0.239 0.336 0.331 BACNE C B 0.440 0.568 0.048 0.012 0.213 0.232 0.354 0.336 0.057 BACNI 0.464 B C 0.301 0.568 0.440 0.048 0.057 0.336 0.239 0.029 B BACPE 0.119 C B 0.060 0.048 0.568 0.415 0.520 0.239 0.336 BACRU C B B 0.440 0.415 0.334 0.048 0.053 B 0.568 0.336 0.255 B BACSA2 C 0.490 0.415 0.568 0.053 0.057 0.240 0.048 0.301 0.336 B BACSA 0.440 C 0.341 0.239 0.415 0.462 0.053 0.048 0.336 0.060 B BACER 0.568 0.119 B C 0.336 0.053 0.560 0.415 0.030 0.295 B BACSP C 0.048 B 0.490 0.720 0.295 0.252 0.053 0.087 B 0.415 C B BAECA 0.295 0.440 0.568 0.415 0.028 0.087 0.336 B 0.053 0.239 B BAESP 0.356 0.295 0.286 0.087 0.380 0.568 0.048 0.053 0.336 C B BAIAL 0.415 0.338 B 0.087 0.295 0.047 0.330 0.565 0.297 0.048 B B BALFO 0.053 C 0.497 0.297 0.087 0.565 0.126 0.336 C 0.295 0.370 B B BALRU 0.297 0.415 0.295 0.470 0.052 0.336 0.126 B 0.087 0.565 B C BALSP 0.297 0.208 0.336 0.370 0.415 0.053 0.087 0.565 0.126 C BAMSP B 0.295 B 0.336 0.297 0.085 0.464 0.409 0.053 0.370 0.126 C B B 0.087 B 0.660 0.340 0.336 0.464 0.119 0.021 0.409 0.297 C B B B 0.565 0.295 0.297 0.085 0.119 0.092 B 0.336 0.464 C B B 0.297 0.126 0.295 0.087 0.336 0.464 0.119 B B B 0.297 C 0.336 0.341 0.359 0.087 0.119 B B C 0.336 B 0.306 0.341 0.119 0.047 0.359 C B B 0.464 0.297 0.336 0.119 0.096 C 0.341 B B 0.119 0.297 0.336 0.341 0.119 C B B 0.356 0.207 0.336 0.119 B C B 0.356 0.338 0.084 0.207 C B 0.341 0.338 0.098 0.356 C B 0.119 0.356 0.338 C B 0.181 0.338 B C 0.336 0.181 C 0.356 0.337 C 0.338 C C . var ca ca fi fi shastensis Appendix C inventory Standing dead tree absolute density and its uncertainty for each decay class by species present in the U.S. FIA GenusAbies Species amabilis Code den0 den1 unc1 cod1 den2 unc2 cod2 den3 unc3 cod3 den4 unc4 cod4 AbiesAbiesAbiesAbies balsamea Abies bracteata Abies concolor Abies fraseri Abies grandis lasiocarpa Abies magni Abies magni AcaciaAcer procera Acer species Acer species Acer barbatum Acer glabrum Acer grandidentatum Acer macrophyllum Acer negundo Acer nigrum Acer pensylvanicum Acer rubrum Acer saccharinum Aesculus saccharum Aesculus species Ailanthus californica spicatum Aleurites species Alnus altissima Alnus fordii Amelanchier rubra species species

22 continued ARMEASTR 0.580BEAL 0.565 0.470BELE 0.126 0.565 0.550BELU 0.600 0.126 0.615 BBENI 0.600 0.673 0.049BEOC B 0.464 0.674 0.049 0.560BEPA B 0.530 0.119 0.464 0.035 0.615BEPA2 B 0.615 0.480 0.119 0.443 0.049 B B 0.480 0.049 0.497 0.462 0.115BEPO B 0.615 B 0.341 0.424 0.032 0.061BESP B B 0.450 0.049 0.119 0.341 0.049 0.443BULA B B 0.615 0.480 0.443 0.119 0.200 B 0.115CADE3 B B 0.049 0.470 0.615 0.115 0.443 0.297 0.084CACA 0.370 B 0.443 B 0.565 0.049 0.356 0.246 0.115 0.087 BCAAQ 0.421 B B 0.580 0.115 0.126 0.338 0.356 0.085 0.278CACO B 0.046 B B 0.610 0.565 0.443 0.278 0.338 0.286 B 0.088CAGL B C B 0.600 0.635 0.126 0.115 0.443 B 0.088 0.370 0.286 0.336CAIL C 0.647 0.278 0.660 0.041 B 0.464 0.115 0.286 0.095 0.337 B BCALA 0.309 B C 0.050 0.088 0.635 0.119 0.600 0.337 B 0.286CAMY B 0.019 B C 0.464 0.278 0.620 0.041 0.286 0.635 B B 0.337CAOV B C 0.560 0.450 0.119 0.088 0.278 0.635 B 0.337 0.286 0.041 BCASP 0.640 0.635 0.404 0.286 C 0.053 0.341 0.088 0.041 0.337 B BCATE 0.328 C B 0.584 0.041 0.052 0.620 0.337 0.450 0.119 BCATO B B 0.058 C 0.044 0.341 0.286 0.540 0.671 0.053 0.450 B B C CADE B 0.640 0.250 0.119 0.337 0.635 0.029 0.286 0.450 B 0.053 B BCAOZ 0.450 0.261 0.400 0.637 0.085 0.041 0.356 0.337 0.053 C B BCAPU 0.262 B 0.400 0.527 0.053 0.085 0.382 0.250 0.041 0.338 BCAST C B B 0.336 0.400 0.382 0.061 0.356 0.459 0.027 0.085 0.250 B B BCATA C 0.291 0.382 0.420 0.087 0.338 0.450 0.045 0.250 C 0.085 B B BCELA 0.250 0.291 0.409 0.337 0.380 0.087 0.565 0.053 0.085 C B BCEOC B 0.323 0.085 0.337 0.383 0.291 0.046 0.470 0.565 0.126 C BCESP B B 0.490 0.383 0.086 0.205 0.046 0.337 0.565 0.126 0.291 C B B B 0.565 0.490 0.383 0.102 0.250 0.084 0.291 0.126 0.337 B B C B 0.291 0.126 0.250 0.565 0.102 0.464 0.085 0.337 B B B C 0.291 0.337 0.268 0.085 0.464 0.126 0.119 B B B C 0.268 0.337 0.291 0.085 0.464 0.119 C B B B 0.464 0.268 0.107 0.291 0.336 0.119 C B B 0.119 0.291 0.464 0.107 0.341 0.337 B C B 0.356 0.337 0.341 0.119 0.119 B B C 0.356 0.338 0.341 0.119 C B B 0.341 0.356 0.338 0.119 C B 0.119 0.341 0.338 0.356 C B 0.356 0.119 0.338 B C 0.356 0.338 B C 0.356 0.338 C 0.338 0.356 C 0.338 C C . var commutata GenusArbutus Species menziesii Code den0 den1 unc1 cod1 den2 unc2 cod2 den3 unc3 cod3 den4 unc4 cod4 Appendix C.—continued AsiminaBetulaBetula triloba BetulaBetula alleghaniensis Betula lenta Betula lutea Betula nigra occidentalis Betula papyrifera Betula papyrifera BumeliaCalocedrus populifolia Carpinus species lanuginosa Carya decurrens Carya caroliniana CaryaCarya aquatica Carya cordiformis Carya glabra Carya illinoensis Carya laciniosa Carya myristicaeformis Carya ovata Castanea species Castanea texana Castanea tomentosa dentata Castanopsis ozarkensis Catalpa pumila species CeltisCeltisCeltis species laevigata occidentalis species

23 continued CECACEIN 0.580CELE 0.565 1.000CEMO 0.126 1.000 0.565CHLA 1.000 0.565 0.126 BCHNO 0.565 0.390 0.126CHTH B 0.420 0.126 0.464 0.377CLLU B 0.377 0.310 0.119 0.041 0.464 BCOFL 0.041 0.377 0.520 0.464 0.119 BCONU B 0.464 0.640 0.041 0.528 0.119 BCOSP B 0.580 0.119 0.341 0.565 0.366 0.125 BCOOB B 0.565 0.366 0.640 0.119 0.126 0.079 0.341 BCRSP B 0.470 0.126 0.079 0.565 0.366 0.341 0.119 BCUSP B B 0.341 0.565 0.620 0.126 0.079 0.485 0.119 B BDIVI B 0.119 0.126 0.440 0.565 0.356 0.464 0.365 0.112 B BEUSP B 0.464 0.365 0.377 0.126 0.338 0.119 0.065 0.356 B B 0.640FAGR B 0.119 0.065 0.464 0.670 0.365 0.041 0.356 0.338 0.565 C BFRAM B B 0.356 0.464 0.560 0.119 0.565 0.065 0.462 0.338 B B B 0.126FRLA C 0.338 0.550 0.119 0.604 0.464 0.126 0.341 0.272 0.102 B BFRNI C 0.341 0.272 0.504 0.366 B 0.038 0.119 0.500 0.119 0.336 C B BFRPE B 0.119 0.336 0.341 0.039 0.272 0.079 0.504 0.450 0.464 B BFRPR C B 0.341 0.119 0.530 0.464 0.336 0.356 0.113 0.504 C B B B 0.119FRQU 0.119 0.540 0.330 0.341 0.504 0.119 0.356 0.338 0.113 B C FRSP B 0.356 0.530 0.315 0.365 B 0.504 0.046 0.119 0.113 0.338 B BGLAQ C B 0.338 0.504 0.356 0.048 0.540 0.065 0.113 0.349 0.341 B BGLTR C B 0.356 0.600 0.113 0.338 0.341 0.504 0.094 0.349 C B B 0.119 BGOLA 0.338 0.565 0.175 0.356 0.600 0.349 0.119 0.113 0.094 C BGYDI B 0.370 0.333 0.272 B 0.349 0.126 0.084 0.338 0.565 0.094 C BHASP B B 0.349 0.565 0.085 0.336 0.094 0.126 0.313 0.500 0.356 C B BILOP B 0.320 0.094 0.126 0.356 0.349 0.114 0.565 0.313 B C 0.338 BJUCI B 0.464 0.403 0.565 0.313 0.338 0.094 0.126 0.500 0.114 B BJUSP B 0.356 C 0.313 0.119 0.336 0.464 0.126 0.114 0.360 0.565 C JUNI B B B 0.313 0.464 0.338 0.114 0.510 0.119 0.356 0.565 0.126 C B B B 0.114 0.119 0.313 0.565 0.338 0.464 0.356 0.510 0.126 C B B B 0.341 0.464 0.356 0.114 0.126 0.119 0.338 0.565 B B C B 0.356 0.119 0.341 0.119 0.338 0.464 0.126 B B B C 0.356 0.341 0.338 0.119 0.464 0.119 B B C B 0.338 0.119 0.356 0.464 0.341 0.119 C B B 0.356 0.341 0.338 0.119 0.119 0.464 C B B 0.338 0.356 0.119 0.341 0.119 C B B 0.356 0.338 0.341 0.119 C B B 0.338 0.341 0.356 0.119 C B 0.356 0.119 0.338 0.341 C B 0.338 0.356 0.119 B C 0.356 0.338 C B 0.356 0.338 C 0.338 0.356 C 0.338 C C orida fl GenusCercis Species canadensis Code den0 den1 unc1 cod1 den2 unc2 cod2 den3 unc3 cod3 den4 unc4 cod4 Appendix C.—continued CercocarpusCercocarpus intricatus Cercocarpus ledifolius Chamaecyparis montanus lawsoniana Chamaecyparis nootkatensis Chamaecyparis thyoides CladrastisCornus Cornus lutea CornusCotinusCrataegus nuttallii Cupressus species Diospyros obovaus species Eucalyptus species Fagus virginiana Fraxinus species FraxinusFraxinus grandifolia americana Fraxinus latifolia Fraxinus nigra Fraxinus pennsylvanica Fraxinus profunda Gleditsia quadrangulata Gleditsia species Gordonia aquatica Gymnocladus triacanthos Halesia dioicus lasianthus IlexJuglansJuglans species Julglans cinerea opaca species nigra

24 continued JUMOJUOC 0.450JUSP2 0.377 0.440LALA 0.440 0.041 0.377LALY 0.528 0.041 0.490 BLAOC 0.125 0.480 0.377 BLASP 0.366 0.480 0.377 0.041 B 0.079 0.377 0.366 0.440 0.041 B 0.485 0.041 0.079 0.377 B B 0.112 0.041 0.366 B B 0.365 0.366 0.079 B B 0.065 0.366 0.365 0.079 B 0.462 0.079 0.065 0.366 B B 0.102 0.079 0.365 B B 0.272 0.365 0.065 B B 0.336 0.365 0.272 0.065 B 0.356 0.065 0.336 0.365 C B 0.338 0.065 0.272 C B 0.272 0.336 C B 0.272 0.336 C 0.336 0.272 C 0.336 C C LISTLITU 0.460LIDE 0.410 0.477MAPO 0.580 0.370 0.022MAAC 0.760 0.565 0.010MASP 0.450 0.565 B 0.126MASP2 A 0.450 0.451 0.126 0.364MEAZ 0.610 B 0.451 0.024 0.350 B 0.045MOAL 0.565 0.470 0.101 0.464 0.010 BMORU 0.126 0.464 B 0.590 0.565 0.119 BMOSP 0.590 A 0.439 0.119 0.565 0.126 B 0.239NYAQ 0.565 B 0.590 0.439 0.055 0.126 0.213 B 0.084 BNYOG 0.464 0.126 0.565 0.460 0.114 0.341 0.084 B BNYSY 0.119 0.460 0.341 B 0.126 0.565 0.464 B 0.119 BOLTE B 0.565 0.263 0.119 0.464 0.460 0.126 0.119 B 0.356 BOSVI 0.464 B 0.126 0.263 0.085 0.119 0.565 1.000 0.237 B 0.338 B BOXAR 0.341 0.119 0.464 0.106 0.126 0.565 0.630 0.356 0.336 B B BPATO 0.119 0.356 C 0.500 0.119 0.464 0.341 0.126 0.565 B 0.338 B BPEBO C 0.464 0.356 0.338 0.555 0.341 0.380 0.119 0.119 B 0.126 BPIAB B 0.341 C 0.119 0.510 0.356 0.338 0.058 0.119 0.565 0.464 C B BPIBR 0.356 B 0.119 0.341 0.565 0.338 0.126 0.119 0.464 0.370 B C B BPIEN 0.338 0.119 0.341 0.356 0.126 0.119 0.330 0.464 0.370 B C B BPIGL2 0.341 0.447 0.356 0.119 0.338 C 0.330 0.389 0.119 0.030 B BPILU B 0.356 0.370 0.119 0.046 0.338 0.464 0.341 0.389 0.024 B C PIMA B A 0.338 0.356 0.389 0.464 0.119 0.119 0.341 0.360 0.024 B C BPIPU B 0.338 0.356 0.024 0.119 0.119 0.380 0.341 0.370 0.380 C B B B 0.356 0.354 0.338 0.389 0.380 0.359 0.119 0.020 0.010 C B B B 0.338 0.086 0.341 0.356 0.024 0.389 0.297 0.017 C B A A 0.359 0.341 0.119 0.338 0.356 0.024 0.013 C B B B 0.017 0.119 0.338 0.356 0.398 0.380 C B B B 0.356 0.359 0.398 0.338 0.065 0.010 B B C 0.338 0.356 0.017 0.359 0.398 0.065 C B A 0.398 0.356 0.338 0.017 0.065 C B B 0.065 0.338 0.260 0.400 C B B 0.398 0.260 0.336 0.020 B C 0.065 0.398 0.197 0.336 C A 0.260 0.065 0.336 B C 0.336 0.254 C B 0.260 0.336 C 0.336 0.260 C 0.336 C C ua fl orus fl GenusJuniperus Species monosperma Code den0 den1 unc1 cod1 den2 unc2 cod2 den3 unc3 cod3 den4 unc4 cod4 JuniperusJuniperusLarix occidentalis Larix species LarixLarix laricina Liquidambar lyallii occidentalis styraci species Appendix C.—continued LiriodendronLithocarpus tulipifera Maclura densi MagnoliaMagnolia pomifera Malus acuminata Melia species MorusMorus species Morus azedarach Nyssa alba Nyssa rubra Nyssa species Olneya aquatica Ostrya ogeche Oxydendrum sylvatica Paulownia tesota arboreum Persea virginiana Picea tomentosa PiceaPicea borbonia Picea abies Picea breweriana Picea engelmannii Picea glauca lutzii mariana pungens

25 continued PIRUPISI 0.380PICE 0.402 0.370PIAL 0.380 0.025 0.406PIAR 0.389 0.370PIAR2 0.010 B 0.370 0.024 0.364PIAT 0.370 B 0.359 0.364 0.015PIBA B 0.364 0.017 0.370 0.015 0.392PIBA2 B 0.015 0.370 0.359 0.364PICL 0.014 B B 0.420 0.364 0.017 0.373 0.015 BPICO 0.390 B 0.449 0.373 0.460 0.015 0.017PICO2 B B 0.373 0.020 0.400 0.067 0.017 0.364 0.373PIDI B 0.370 B 0.017 0.390 0.398 0.373 0.015 APIEC 0.057 B B 0.364 0.001 0.373 0.065 0.388 0.017 B 0.500PIED B 0.015 0.380 B 0.470 0.260 0.388 0.017 0.060 0.364PIEL A B B 0.388 0.010 0.500 0.364 0.336 0.060 0.373 B 0.015 0.323PIEN2 B B 0.060 0.415 0.364 0.260 0.015 0.540 0.388 0.017 APIFL 0.336 C B 0.373 0.370 B 0.015 0.388 0.015 0.336 0.364 0.284 0.060 BPIGL B B 0.017 0.364 0.320 C 0.284 0.060 0.370 0.015 0.336 0.373PIJE A C B B 0.284 0.015 0.040 0.410 0.373 0.336 0.388 B 0.364 0.017PILA B C B 0.336 0.400 0.373 0.389 0.017 0.284 0.370 0.060 0.015 B APILE C 0.388 B 0.001 0.017 0.284 0.340 0.021 0.373 0.336 0.361 C PIMO B B 0.060 B 0.373 0.284 0.370 0.336 0.365 0.017 0.014 0.388PIMO3 A B B C 0.017 0.336 0.500 0.388 0.284 0.364 B 0.373 0.017 0.060PIMU 0.350 C B B 0.364 0.284 0.388 0.359 0.060 0.336 0.015 0.017 B C PINI 0.341 0.284 B B 0.370 0.015 0.336 0.060 0.017 0.388 0.412PIPA B 0.010 C B 0.336 B 0.398 0.364 0.307 0.060 0.019 0.410 0.284PIPO B C B B 0.060 0.015 0.540 0.284 B 0.373 C 0.388 0.012 0.364 0.336PIPU2 B B 0.373 0.380 0.284 0.364 0.398 0.336 0.017 0.060 B 0.015PIRA B 0.348 B 0.490 C 0.017 0.335 0.336 0.015 0.060 0.284 0.308PIRE C 0.012 B B 0.389 0.284 B 0.373 0.370 0.010 0.220 0.336 0.058PIRI B C B B 0.021 0.336 0.017 0.410 0.364 B 0.388 0.284 0.057 0.373PISA B C B 0.388 0.337 0.373 0.015 0.260 0.060 C 0.336 B 0.470 0.017 B 0.437 B 0.060 0.383 0.370 0.019 0.017 0.336 0.344 0.364 B 0.057 B C 0.359 B 0.388 0.013 0.364 0.205 0.336 0.015 B B C B 0.017 0.060 0.373 B 0.015 0.284 0.336 0.388 B C B 0.284 0.373 0.388 0.017 0.336 B 0.060 B 0.284 B C 0.336 0.469 0.017 0.060 0.373 B 0.336 C 0.398 B 0.284 0.058 0.373 0.017 C B B 0.060 0.336 0.388 C 0.017 0.284 B B 0.383 0.284 0.060 B 0.336 C B 0.217 0.060 0.336 0.388 B 0.260 C 0.336 0.388 0.060 B C 0.336 0.284 0.060 C B 0.252 0.336 C B 0.336 0.284 C 0.284 0.336 C 0.336 C C exilis fl GenusPicea Species rubens Code den0 den1 unc1 cod1 den2 unc2 cod2 den3 unc3 cod3 den4 unc4 cod4 Appendix C.—continued PiceaPiceaPinusPinus sitchensis Pinus species Pinus albicaulis Pinus aristata Pinus arizonica Pinus attenuata Pinus balfouriana Pinus banksiana Pinus clausa Pinus contorta Pinus coulteri Pinus discolor Pinus echinata Pinus edulis Pinus elliotti Pinus engelmannii PinusPinus glabra Pinus jeffreyi Pinus lambertiana Pinus leiophylla Pinus monophylla Pinus monticola Pinus muricata Pinus nigra Pinus palustris Pinus ponderosa Pinus pungens Pinus radiata resinosa rigida sabiniana

26 continued PISEPIST2 0.510PIST 0.350 0.364PISY 0.364 0.340 0.015PITA 0.015 0.380 0.364PIVI B 0.470 0.370 0.015 BPLAQ 0.373 0.382 0.010 0.450PLOC B 0.373 0.530 0.017 0.011 0.364POAL A 0.460 0.017 0.565 0.373POAN 0.015 B B 0.370 0.565 0.126 0.360 0.017 BPOBA 0.340 B 0.374 0.126 0.388 0.352 0.010PODE B B 0.310 0.374 0.388 0.085 0.060 0.012 0.373 BPOFR A 0.370 0.374 0.085 0.060 0.464 0.388POGR 0.017 B B B 0.340 0.374 0.464 0.085 0.119 0.310 0.060 B BPOHE 0.360 0.374 0.085 B 0.464 0.119 0.284 0.325 0.030 BPOSA B B 0.356 0.370 0.464 0.284 0.085 0.119 0.336 0.057 B 0.388 BPOSP A 0.031 0.370 0.374 0.464 0.119 0.336 0.341 0.284 BPOTR 0.060 B C B 0.370 0.464 0.374 0.085 0.341 0.119 0.119 0.284 0.336 B B C PRSP 0.390 0.392 0.464 0.119 0.085 B 0.314 0.119 0.324 0.336 B BPRAM B C 0.464 0.314 0.580 0.370 0.033 0.119 0.115 0.336 B B 0.284 BPRNI C 0.470 0.119 0.464 0.314 0.115 0.565 0.020 0.356 B BPRPE 0.336 B C 0.518 0.314 0.464 0.119 0.356 0.115 0.126 0.338 0.470 B B APRSE 0.044 0.360 0.464 0.314 0.115 0.119 C 0.269 0.338 0.518 B B BPRSP2 C 0.313 0.269 0.470 0.290 0.425 0.119 0.115 0.337 0.044 B B B C PRVI 0.470 0.086 0.314 0.269 0.337 0.612 0.464 0.020 0.024 B BPSME C 0.518 B 0.460 0.269 0.314 0.115 0.337 0.047 0.119 0.360 B C A BQUAG 0.044 0.450 0.052 0.315 0.269 0.337 0.115 0.460 0.518 B C B BQUAL 0.700 0.269 0.382 0.210 0.371 0.086 0.337 B 0.052 0.044 B C BQUAR 0.648 0.337 0.269 0.600 0.011 0.549 0.341 0.030 0.045 C BQUBI 0.460 B B 0.700 0.056 0.295 0.269 0.337 0.601 0.050 0.119 C B A BQUCH 0.052 0.648 0.112 0.269 0.337 0.028 0.640 0.295 0.460 B C B BQUCO 0.700 0.354 0.056 0.269 0.227 0.336 0.648 B 0.112 0.052 B C BQUDO 0.600 0.648 0.495 0.013 0.363 0.356 0.337 0.084 0.056 B C 0.295 B B 0.510 0.605 0.056 0.062 0.356 0.426 0.090 0.338 B C B B 0.112 0.648 0.037 0.495 0.338 0.045 0.356 0.295 B B C B 0.056 0.216 0.062 0.356 0.495 B 0.338 0.112 B C B 0.495 0.401 0.057 0.356 0.338 0.062 B B 0.356 C B 0.552 0.062 0.092 0.352 0.338 B C B 0.338 0.495 0.049 0.401 0.085 0.356 B B C 0.062 0.207 0.092 0.401 C 0.338 B B 0.401 0.405 0.336 0.092 B B C 0.464 0.092 0.336 0.282 C B 0.401 0.085 0.405 0.336 C B 0.092 0.336 0.405 B C 0.405 0.336 B C 0.405 0.336 C 0.405 0.336 C 0.336 C C GenusPinus Species serotina Code den0 den1 unc1 cod1 den2 unc2 cod2 den3 unc3 cod3 den4 unc4 cod4 Appendix C.—continued PinusPinusPinusPinus strobiformis Pinus strobus Planera sylvestris Platanus taeda Populus virginiana Populus aquatica occidentalis Populus alba Populus angustifolia Populus balsamifera Populus deltoides Populus fremontii Populus grandidentata Populus heterophylla Populus sargentii Prosopis species Prunus tremuloides Prunus species PrunusPrunus americana Prunus nigra Prunus pensylvanica Pseudotsuga serotina Quercus species menziesii Quercus virginiana Quercus agrifolia Quercus alba Quercus arizonica, grisea Quercus bicolor Quercus chrysolepis coccinea douglasii

27 continued QUDUQUEL 0.600QUEM 0.648 0.560QUEN 0.700 0.056 0.648QUFA 0.648 0.700 0.056 BQUFA2 0.056 0.648 0.520 B 0.610 0.495 0.056 0.648QUGA B 0.648 0.062 0.495 0.056QUGA2 B 0.640 0.056 0.495 0.062 0.640QUHY B B 0.648 0.062 0.495 B 0.648QUIL B 0.700 0.056 0.401 0.062 0.495QUIM 0.056 B 0.495 0.648 0.092 0.560 0.401 0.062 BQUIN B 0.062 0.056 0.560 B 0.401 0.648 0.092QUKE B B 0.495 0.648 0.560 0.092 0.401 0.056 B B 0.495QULA B 0.510 0.062 0.405 0.056 0.648 0.092 0.401QULA2 0.062 B B 0.401 0.495 0.648 0.520 0.336 0.056 0.405 0.092 BQULO B 0.560 B 0.092 0.062 0.056 0.648 B 0.405 0.495 0.336QULY C B 0.648 B 0.401 0.640 0.056 0.495 0.336 0.405 0.062 B B BQUMA 0.401 0.056 C 0.092 0.648 0.570 0.062 0.495 0.336 0.405 BQUMA2 C 0.092 B 0.580 0.405 0.401 0.495 0.056 0.648 0.062 B 0.336 B 0.560QUMI B C 0.648 0.336 0.092 0.062 0.495 0.056 B 0.401 B 0.648QUMU B 0.495 C 0.056 0.405 0.600 0.062 0.401 0.092 C B B BQUNI 0.056 0.405 0.062 0.600 0.336 0.495 0.648 0.092 0.401 B BQUNU 0.336 B 0.648 0.405 0.401 0.062 0.495 B 0.560 0.056 0.092 B C QUOB B 0.560 0.495 0.056 0.336 0.092 0.401 0.062 C 0.648 0.405 B 0.495QUPA B B 0.401 0.700 0.648 0.062 0.092 0.405 0.056 0.336 B C B BQUPH 0.062 0.092 0.648 0.580 0.056 0.401 0.495 0.336 0.405 B BQUPR B C 0.495 0.560 0.056 0.648 0.405 0.092 0.401 B 0.062 0.336 B BQURU C 0.401 0.062 0.570 0.648 0.056 0.336 0.405 0.092 0.495 B B 0.401QUSH B C 0.405 0.560 0.495 0.092 0.773 0.056 0.336 0.062 B C B BQUSP 0.092 0.336 0.560 0.578 0.495 0.062 0.061 0.405 0.401 B B C QUST B 0.401 0.580 0.648 0.032 0.062 0.495 0.336 0.405 B 0.092 C B BQUVE 0.405 0.092 0.495 0.600 0.599 0.056 0.062 0.336 0.401 B B C 0.405 B 0.401 0.336 0.560 0.538 0.062 0.648 0.024 0.092 B B B C 0.336 0.333 0.401 0.092 0.648 0.063 0.056 0.405 C B B B 0.405 0.495 0.051 0.092 0.401 0.056 C 0.336 B B B 0.336 0.401 0.508 0.062 0.092 0.405 B B B C 0.405 0.309 0.092 0.495 0.048 0.336 C B B 0.406 0.405 0.336 0.399 0.085 0.062 B B C 0.401 0.090 0.336 0.405 0.056 C B B 0.405 0.334 0.092 0.336 C B B 0.405 0.336 0.401 0.085 B C 0.393 0.476 0.336 0.092 C B 0.405 0.336 0.095 C B 0.329 0.336 C B 0.405 0.336 C 0.405 0.336 C 0.336 C C . falcata . var var pagodaefolia GenusQuercus Species durandii Code den0 den1 unc1 cod1 den2 unc2 cod2 den3 unc3 cod3 den4 unc4 cod4 Appendix C.—continued QuercusQuercusQuercus ellipsoidalis Quercus emoryi Quercus engelmannii Quercus falcata Quercus falcata Quercus gambelii Quercus garryana Quercus hypoleucoides Quercus ilicifolia Quercus imbricaria Quercus incana Quercus kelloggii Quercus laevis Quercus laurifolia Quercus lobata Quercus lyrata Quercus macrocarpa Quercus marilandica Quercus michauxii Quercus muehlenbergii Quercus nigra Quercus nuttalli Quercus oblongifolia Quercus palustris Quercus phellos Quercus prinus Quercus rubra Quercus shumardii Quercus species stellata velutina

28 continued QUVIQUWI 0.800RONE 0.700 0.648ROPS 0.660 0.648 0.056SASP 0.660 0.769 0.056SASE B 0.769 0.170 0.360 BSAAL 0.031 0.470 0.565 0.495 BSESE 0.495 0.565 0.420 0.126 0.062 BSEGI 0.616 0.340 0.062 0.126 0.458SOAM B B 0.616 0.154 0.377 0.024 0.340 BTASP B 0.420 0.053 0.464 0.041 0.401 0.377 BTADI B 0.565 0.401 0.400 0.464 0.119 0.092 0.041 B B 0.126 0.341 0.092 0.565 0.119 0.427 0.420TABR B B B 0.341 0.119 0.366 0.126 0.047 0.377THOC B B B 0.600 0.119 0.341 0.079 0.405 0.366 0.041THPL B B B 0.290 0.377 0.464 0.405 0.341 0.119 0.336 0.079TIAM B B 0.326 B 0.310 0.041 0.119 0.356 0.336 0.464 0.119 0.355TIHE B C B 0.016 0.315 0.320 0.356 0.338 0.365 0.119 0.084 0.366TISP B B C B 0.016 0.430 0.338 0.320 0.356 0.065 0.365 B 0.079TOCA C B B 0.366 0.341 0.023 0.320 0.356 0.430 0.338 0.065TSCA B C B 0.340 0.298 B 0.079 0.119 0.341 0.430 0.338 0.097 0.356TSHE B C B 0.377 0.102 0.400 0.298 0.272 0.119 0.097 0.338 0.365TSME B B C B 0.440 0.041 0.394 0.102 0.366 0.336 0.272 B 0.065TSSP B B C 0.420 0.390 0.365 0.025 0.356 0.045 0.366 0.336 BULAL B C 0.447 0.376 B 0.380 0.010 0.065 0.338 0.356 0.366 0.098ULAM B B C 0.366 0.059 0.025 0.376 0.570 0.352 0.338 0.098 0.272ULCR A B C B 0.460 0.079 0.370 0.025 0.565 0.057 0.269 B BULPU 0.336 C B 0.570 0.565 0.360 0.272 0.080 0.126 0.084 0.269 BULRU B B 0.240 0.370 0.460 0.565 C 0.126 0.020 0.336 0.269 0.107ULSE B B B 0.480 0.365 0.336 0.080 0.565 0.126 0.370 0.222 0.107 BULSP C A B 0.565 0.065 0.570 0.462 0.126 0.080 0.464 0.336 0.356 C B BULTH B 0.464 0.500 0.126 0.290 0.565 0.063 0.119 0.338 0.356 BUMCA B B C 0.400 0.464 0.119 0.570 0.565 0.020 0.126 0.356 0.338 BVAAR B B C 0.510 0.272 0.066 0.464 0.119 0.565 0.400 0.126 0.338 B A B C 0.565 0.470 0.464 0.336 0.299 0.119 0.126 0.066 0.341 B B B C 0.126 0.341 0.565 0.119 0.170 0.464 0.336 0.119 C B B B 0.299 0.341 0.119 0.126 0.464 0.024 0.119 B B C B 0.336 0.341 0.119 0.464 0.299 0.119 B B A B 0.464 0.341 0.119 0.119 0.336 0.356 C B B 0.119 0.356 0.464 0.119 0.341 0.338 B C B 0.356 0.338 0.119 0.341 0.119 B B C 0.356 0.338 0.341 0.119 C B B 0.341 0.356 0.338 0.119 C B 0.119 0.341 0.338 0.356 C B 0.119 0.356 0.338 B C 0.356 0.338 B C 0.356 0.338 C 0.338 0.356 C 0.338 C C . var nutans GenusQuercus Species virginiana Code den0 den1 unc1 cod1 den2 unc2 cod2 den3 unc3 cod3 den4 unc4 cod4 Appendix C.—continued QuercusRobiniaRobinia wislizeni SalixSapium neomexicana Sassafras pseudoacacia Sequoia species Sequoiadendron sebiferum albidum giganteum Sorbus sempervirens TamarixTaxodium americana Taxus species distichum ThujaThujaTilia brevifolia Tilia occidentalis Tilia plicata Torreya americana Tsuga heterophylla Tsuga species Tsuga californica Tsuga canadensis Ulmus heterophylla Ulmus mertensiana Ulmus species Ulmus alata Ulmus americana Ulmus crassifolia Ulmus pumila Ulmus rubra Umbellularia serotina Vaccinium species californica thomasii arboreum

29 3 3 3 3 3 nition Code/Units Method fi Appendix C.—continued Standing Dead Absolute Density Predictions Metadata (Appendix C) Fieldnamegenusspecies De codeden0 Sample genus Sample species den1unc1 Genus-species code cod1 Green density (decay class “0”)cod1cod1 Density of decay class 1 snag Uncertainty of den1 den2 Uncertainty code for den1 (A, B, C)unc2 Uncertainty code for den1 (A, B, C) g/cm cod2 Uncertainty code for den1 (A, B, C)cod2 g/cm cod2 Density of decay class 2 snag A Uncertainty of den2 Bden3 Uncertainty code for den2 (A, B, C) Cunc3 Uncertainty code for den2 (A, B, C)cod3 Uncertainty code for den2 (A, B, C) Species sampled cod3 g/cm Genera sampled cod3 Density of decay class 3 snag A Species and genus not sampled Uncertainty of den3 Bden4 Uncertainty code for den3 (A, B, C) Cunc4 Uncertainty code for den3 (A, B, C)cod4 Uncertainty code for den3 (A, B, C) Species sampled cod4 g/cm Genera sampled cod4 Density of decay class 4 snag A Species and genus not sampled Uncertainty of den4 B Uncertainty code for den4 (A, B, C) C Uncertainty code for den4 (A, B, C) Uncertainty code for den4 (A, B, C) Species sampled g/cm Genera sampled A Species and genus not sampled B C Species sampled Genera sampled Species and genus not sampled

30 continued ABAMABBA 0.400ABBR 0.936 0.340ABCO 0.044 1.040 0.360ABFR 0.062 1.018 0.370 BABGR 0.056 0.996 0.340 B 0.955ABLA 0.021 0.350 1.018 B 0.022 1.150ABMA 1.013 0.056 0.310 B 0.023 0.975 B 0.011 0.360 1.040 BABMA2 0.091 0.873 B 1.040 0.030 0.803 B 0.360ABPR 0.018 0.975 0.017 B 0.107 1.018 1.101 BABIE 0.966 0.091 0.370 B 0.056 0.064 B 0.803ACSP2 B 0.021 1.035 1.068 0.340 B 0.107 0.625 0.600ACBA B B 0.013 1.080 0.073 0.574 B 1.018 0.054 0.803 0.982ACGL 0.051 B 0.540 0.216 0.056 0.975 0.710 B B 0.855 0.107 0.129ACGR B 0.979 0.440 0.091 0.219 B 0.574 C 0.054 B 0.836 1.000ACMA B 0.035 B 0.440 0.979 0.216 0.541 C B 0.013 0.626 0.056ACNE B 0.975 0.979 0.035 0.440 0.213 0.574 B 0.793 C 0.054 0.091ACNI 0.035 0.803 B 0.979 B 0.440 0.574 0.216 0.201 B C 0.766ACPE 0.107 B 0.035 0.979 0.216 B B 0.845 0.520 0.696 C 0.194 B 0.766ACRU 0.035 0.440 B 0.054 0.467 0.979 B 0.214 C 0.803 0.766 0.194ACSA2 B 0.618 0.979 0.490 0.213 0.035 B 0.107 0.194 0.574 B 0.766 C 0.440ACSA 0.171 B 0.035 1.002 0.565 0.216 C B 0.194 0.766 0.979 BACER B 0.575 0.049 0.560 0.160 B 0.565 B 0.194 0.035 C 0.213ACSP 0.766 B 0.956 0.490 0.622 0.565 0.160 B B 0.525 0.766 0.194AECA B 0.028 0.979 B 0.217 0.160 0.440 C 0.565 0.223 B 0.194 0.773AESP 0.450 0.035 0.979 B 0.160 0.380 0.565 B 0.766 C B 0.056 0.221 C AIAL 0.450 B 0.035 0.982 0.330 0.160 B 0.194 0.565 B 0.761 0.450 0.221ALFO B 0.129 0.982 C 0.565 B 0.160 0.370 B 0.055 0.766 B 0.221 0.450 0.129 C 0.470 0.160 0.618 B 0.982 0.194 0.766 B 0.221 0.450 B 0.565 C 0.982 0.086 0.129 B B 0.194 0.793 0.221 B 0.160 C 0.129 0.450 0.526 B 0.201 B 0.793 0.450 B 0.221 C 0.077 0.565 B B 0.201 0.221 0.450 B 0.793 0.160 0.565 C B 0.450 0.793 0.221 0.201 B C 0.160 0.618 B 0.221 0.201 0.450 C 0.171 B 0.618 B 0.213 C 0.450 B 0.171 B 0.618 0.221 0.450 C 0.618 0.171 B 0.221 0.525 C 0.171 0.223 B 0.525 C B 0.223 C 0.525 0.525 0.223 C 0.223 C C . var ca ca fi fi Speciesamabilis balsamea Codebracteata concolor den0fraseri rel1grandis unc1lasiocarpa cod1magni rel2shastensis unc2procera cod2species rel3species barbatum unc3glabrum cod3grandidentatum rel4macrophyllum unc4negundo cod4 nigrum pensylvanicum rubrum saccharinum saccharum species spicatum californica species altissima fordii Appendix D inventory Standing dead tree relative density and its uncertainty for each decay class by species present in the U.S. FIA Genus Abies Abies Abies Abies Abies Abies Abies Abies AbiesAbies Abies magni Acacia Acer Acer Acer Acer Acer Acer Acer Acer Acer Acer Acer Acer Aesculus Aesculus Ailanthus Aleurites

31 continued ALRUALSP 0.370AMSP 1.030 0.370ARME 0.013 0.660 1.030ASTR 0.982 0.580 0.135 BBEAL 0.129 0.982 0.470 B 0.903BELE 0.129 0.982 0.550 B 0.051 0.903BELU 0.129 1.032 0.600 B 0.793 0.228BENI B 0.135 1.040 0.600 B 0.201 0.793BEOC B 0.070 1.040 0.535 0.560 B 0.201 0.793BEPA B 0.044 0.530 0.074 1.032 0.535 B 0.201 0.713 B 1.032 0.480 0.135 0.618 0.153 BBEPA2 B 0.181 0.756 0.135 B 1.016 0.171 0.618BEPO 0.480 B B 0.077 0.671 0.393 0.052 B 0.171 B 0.618 1.032BESP B 0.048 0.450 0.213 0.713 0.393 B 0.171 0.135 B 0.336BULA B 0.713 1.032 0.480 0.181 0.525 0.219 B C 0.075 0.500CADE3 0.181 0.135 B B 1.032 0.713 0.223 0.470 0.525 B C 0.084 0.400 0.370CACA 0.135 0.181 B 0.982 0.223 B B 0.525 0.713 C 0.076 1.040CAAQ 0.511 B 0.129 0.580 0.223 0.181 B B 0.402 C 0.511 0.713 0.074 0.148CACO B 0.982 0.610 0.213 0.439 B 0.148 C 0.181 B 0.713 0.777CAGL 0.129 0.988 B 0.600 B 0.220 0.439 C 0.181 0.120 0.793CAIL B B 0.069 1.040 0.511 0.660 0.220 B 0.972 0.439 C 0.201CALA 0.075 0.148 B B 0.988 0.439 0.511 B 0.035 0.600 0.220 C 0.793CAMY 0.069 B 0.220 0.148 0.620 B B 0.511 0.439 0.988 0.201 0.705 B C CAOV 0.560 0.988 0.148 0.220 0.069 B 0.618 C B 0.064 0.661 0.439CASP B 1.011 0.988 0.069 0.640 0.171 0.056 0.220 C B B 0.705CATE 0.439 B 0.059 0.069 0.895 0.620 0.618 B 0.064 B 0.220CATO B C 0.439 0.062 0.705 1.040 0.540 0.171 0.404 B B 0.705CADE 0.220 0.064 B 0.525 0.033 0.988 0.640 C 0.080 0.440 B B 0.596 0.705 0.064CAOZ 0.223 0.069 0.977 0.400 0.077 C B 0.404 B B 0.218 0.064 0.808 0.525 0.056 B 0.936 0.400 C 0.080 B B 0.073 0.404 0.727 0.223 C 0.396 B 0.043 0.936 B 0.404 0.080 B 0.035 0.705 0.219 0.396 B 0.123 C 0.404 0.080 B 0.064 0.627 0.219 B 0.396 B C 0.080 0.510 B 0.038 B 0.940 0.219 B C 0.081 0.396 0.334 B 0.046 0.940 B 0.396 0.219 C 0.074 0.404 B 0.237 0.396 0.219 B 0.080 0.332 C B 0.219 0.396 B 0.078 C 0.668 B 0.219 0.396 C 0.082 0.676 B 0.213 0.396 C 0.185 B 0.219 0.396 C B 0.219 0.525 C 0.223 0.525 C 0.223 C C Speciesrubra species Codespecies menziesii den0triloba rel1alleghaniensis unc1lenta cod1lutea rel2nigra occidentalis unc2papyrifera cod2 rel3commutata populifolia unc3species cod3lanuginosa rel4decurrens unc4caroliniana cod4 aquatica cordiformis glabra illinoensis laciniosa myristicaeformis ovata species texana tomentosa dentata ozarkensis Genus Alnus Appendix D.—continued Alnus Amelanchier Arbutus Asimina Betula Betula Betula Betula Betula Betula BetulaBetula Betula papyrifera var. Bumelia Calocedrus Carpinus Carya Carya Carya Carya Carya Carya Carya Carya Carya Carya Castanea Castanea

32 continued CAPUCAST 0.400CATA 0.936 0.420CELA 0.123 0.982 0.380CEOC 0.129 0.982 0.470 BCESP 0.129 0.490 0.982 B 0.940CECA 0.982 0.129 0.490 B 0.237 0.793CEIN 0.129 0.982 0.580 B 0.201 0.793CELE B 0.129 0.982 B 1.000 0.201 0.793CEMO 0.129 B 1.000 0.982 0.676 B 0.793 0.201CHLA B 1.000 0.982 0.129 0.185 B 0.618 0.201 0.793CHNO 0.982 B 0.129 0.390 0.171 0.618 B B 0.201 0.793CHTH 0.129 B 0.420 0.994 0.171 0.618 B 0.201 BCLLU 0.793 0.525 B 0.994 0.071 0.310 B 0.618 0.171 B 0.793 0.201 0.223COFL 0.071 B 0.525 0.994 0.171 0.520 0.618 B 0.793 B 0.201CONU 0.223 0.525 0.071 C B 0.982 0.640 0.171 B 0.618 0.201 B 0.951COSP 0.223 0.525 B 0.580 0.129 0.982 C 0.171 B 0.618 B 0.951 0.260 B 0.525COOB 0.223 0.982 0.129 0.640 C 0.618 0.171 B 0.260 B 0.951 0.223CRSP 0.129 0.525 B 0.982 0.470 0.618 C 0.171 B 0.260 B 0.844 0.223 BCUSP 0.525 0.129 0.982 C 0.171 0.620 B 0.902 B 0.199 0.793 0.223DIVI 0.129 B 0.982 0.525 0.440 C 0.902 0.146 B B 0.793 0.201EUSP 0.525 0.129 0.223 B 0.994 0.146 C B 0.902 0.201 0.640 B 0.793 0.525FAGR 0.223 0.071 B 0.670 0.146 B C 0.775 B 0.982 0.201 0.793 0.223 BFRAM 0.605 0.982 0.560 C 0.163 B 0.618 0.129 0.201 B 0.793 0.605FRLA 0.218 B C 0.129 1.015 0.550 0.618 0.171 0.201 B 0.951 0.218 B BFRNI 0.605 0.056 0.899 0.171 C 0.618 0.500 B 0.260 B 0.218FRPE B 0.061 0.525 C 0.793 0.171 0.899 0.618 B 0.450 B 0.793FRPR 0.223 B 0.525 0.201 0.118 0.171 C 0.530 0.618 0.899 B B 0.201 0.663 0.525FRQU 0.223 0.899 0.540 0.171 0.118 C 0.902 B B B 0.041 0.562 0.223 0.525FRSP B 0.118 0.899 0.530 C 0.146 B B 0.049 0.618 0.223 0.622 0.525 B 0.118 0.899 C 0.540 0.618 B B 0.171 0.159 0.223 0.525 0.622 B 0.118 C 0.899 0.171 0.508 B 0.622 0.223 0.159 0.605 B C 0.118 B 0.076 0.604 B B 0.159 0.622 0.218 C B 0.079 0.525 B 0.575 B 0.159 0.622 0.525 B C 0.223 0.162 0.575 B 0.159 0.622 0.223 0.533 B 0.575 0.162 C 0.159 B 0.213 0.525 B C 0.162 0.575 B 0.223 B 0.525 C 0.162 0.575 B 0.223 C 0.525 0.162 0.575 B 0.525 0.223 C 0.162 B 0.223 0.525 C B 0.223 0.525 C 0.223 0.525 C 0.223 C C orida Speciespumila species Codespecies laevigata den0occidentalis rel1species unc1canadensis cod1intricatus rel2ledifolius montanus unc2lawsoniana cod2nootkatensis rel3thyoides unc3lutea cod3fl rel4nuttallii species unc4obovaus cod4 species species virginiana species grandifolia americana latifolia nigra pennsylvanica profunda quadrangulata species Genus Castanea Appendix D.—continued Castanopsis Catalpa Celtis Celtis Celtis Cercis Cercocarpus Cercocarpus Cercocarpus Chamaecyparis Chamaecyparis Chamaecyparis Cladrastis Cornus Cornus Cornus Cotinus Crataegus Cupressus Diospyros Eucalyptus Fagus Fraxinus Fraxinus Fraxinus Fraxinus Fraxinus Fraxinus Fraxinus

33 continued LIST 0.460 1.017 0.013 B 0.778 0.039 B 0.526 0.075 B 0.525 0.223 C LITULIDE 0.410MAPO 0.910 0.580MAAC 0.760 0.030 0.982MASP 0.982 0.450 0.129 AMASP2 0.129 0.884 0.450 B 0.610MEAZ 0.026 0.850 0.884 B 0.982MOAL 0.020 0.116 0.793 0.470 B 0.793 0.129MORU 0.201 0.982 0.590 A B 0.201 0.862MOSP 0.590 0.129 B 0.982 B 0.074 0.554 0.862 BNYAQ 0.982 0.129 0.590 B 0.793 0.075 0.218 0.618 0.129NYOG B 0.982 0.460 0.618 0.201 B 0.171 0.793NYSY B 0.129 B 0.982 0.460 0.171 B 0.530 0.201 B 0.793 B 0.129 0.982 0.080 0.460 0.501 B 0.530 B 0.793 0.201 0.129 B 0.618 0.982 0.214 0.152 0.525 B 0.201 B 0.793 0.525 0.171 B 0.129 0.223 B 0.618 C 0.201 B 0.793 0.223 B 0.525 0.171 B 0.618 B C 0.201 0.793 0.223 B 0.525 C 0.618 0.171 0.201 B 0.525 0.793 0.223 B 0.171 C 0.618 0.223 B 0.201 B 0.525 C 0.171 0.618 B C 0.223 0.525 B 0.171 0.618 B 0.525 0.223 0.171 C 0.618 B 0.223 0.525 C 0.171 B 0.223 0.525 C B 0.223 0.525 C 0.223 0.525 C 0.223 C C GLAQGLTR 0.600GOLA 0.982 0.600GYDI 0.129 0.982 0.370HASP 0.129 0.982 0.500 BILOP 0.129 0.320 0.982 B 0.793JUCI 0.982 0.129 B 0.500 0.201 0.793JUSP 0.129 0.982 0.360 B 0.201 0.793JUNI B 0.510 0.129 B 0.982 0.201 0.793JUMO B 0.982 0.618 0.129 0.510 B 0.793 0.201JUOC B 0.450 0.129 0.171 0.618 0.982 B 0.201JUSP2 0.793 0.994 B 0.440 0.171 0.618 0.129 B B 0.201LALA 0.440 B 0.071 0.793 0.994 0.171 0.618 B B 0.793LALY 0.525 0.982 0.201 0.071 B B 0.618 0.490 0.171 B 0.201 0.223 0.129LAOC 0.525 0.793 B 0.171 0.994 0.480 B 0.618 0.951 BLASP 0.223 0.525 0.201 B C B 0.480 0.071 0.994 0.171 B 0.260 0.618 0.951 0.223 0.525 0.994 0.071 C B 0.440 0.618 0.844 0.171 B 0.260 B B 0.525 0.223 0.071 C 0.994 0.171 0.199 B 0.618 B 0.223 0.951 B 0.071 0.525 0.902 C B 0.171 B B 0.260 0.951 0.223 C 0.146 0.525 0.902 B 0.951 0.260 B 0.525 0.775 0.223 B 0.146 C B 0.260 0.951 0.223 0.163 B 0.525 C 0.902 B 0.260 0.605 B 0.223 C B 0.146 0.902 0.218 0.605 B 0.902 0.146 C 0.525 B 0.218 C 0.146 0.902 0.223 B 0.605 C 0.146 B C 0.218 0.605 B 0.605 0.218 C 0.218 0.605 C 0.218 C C ua fl orus fl tulipifera densi pomifera acuminata species species azedarach alba rubra species aquatica ogeche sylvatica Speciesaquatica triacanthos Codelasianthus dioicus den0species rel1opaca unc1cinerea cod1species rel2nigra monosperma unc2occidentalis cod2species rel3laricina unc3lyallii cod3occidentalis rel4species styraci unc4 cod4 Liriodendron Appendix D.—continued Lithocarpus Maclura Magnolia Magnolia Malus Melia Morus Morus Morus Nyssa Nyssa Nyssa Genus Gleditsia Gleditsia Gordonia Gymnocladus Halesia Ilex Juglans Juglans Julglans Juniperus Juniperus Juniperus Larix Larix Larix Larix Liquidambar

34 continued OLTEOSVI 1.000OXAR 0.982 0.630PATO 0.129 0.500 0.982PEBO 1.040 0.380 0.129 BPIAB 0.104 0.982 0.510 B 0.793PIBR 0.129 0.982 B 0.370 0.201PIEN 0.793 0.129 1.010 B 0.330 0.877 0.201PIGL2 B 0.083 1.038 B 0.330 0.043 0.793PILU 0.370 B 0.074 0.953 0.618 A 0.201 0.793PIMA B 1.038 0.068 0.171 0.360 0.618 B 0.201PIPU 1.010 0.074 B 0.714 0.380 1.060 0.171 B B 0.040PIRU 0.987 B 0.084 1.038 0.020 B 0.618 0.380 B 0.270PISI 0.899 0.525 0.074 A 0.171 0.618 1.038 0.380 B A 0.987 0.021PICE 0.223 0.525 B 0.171 0.074 1.040 B 0.991 0.270 B 0.370 0.525PIAL 1.060 0.223 B 0.058 C 0.380 0.159 0.991 1.040 B B 0.223 0.987PIAR 0.020 B 0.525 1.038 C 0.159 0.370 0.013 0.991 B 0.270 B 0.223PIAR2 0.525 0.987 C 0.074 A 0.991 0.370 0.953 0.159 B B 0.223 0.270PIAT 0.987 0.370 B 0.605 C 0.159 0.953 0.057 B 1.110 B 0.270PIBA 0.953 0.218 0.605 C 0.998 B 0.057 0.370 0.991 0.040 B BPIBA2 0.987 0.057 0.218 0.022 0.504 B 0.953 0.159 0.370 C 0.991 B 0.270 A 0.605PICL 0.950 0.213 0.420 B 0.057 0.916 C B 0.159 1.107 B 0.218PICO 0.950 0.075 0.953 B 0.564 0.061 C 0.107 0.460 0.950 B 0.984 B 0.075PICO2 0.040 0.605 C 0.213 B 0.400 0.953 0.991 0.075 A 0.055 BPIDI 0.950 0.218 0.370 0.605 B 0.980 0.057 A C 0.159 0.927PIEC B 0.075 0.876 0.953 B 0.218 0.605 0.010 C 0.927 0.091 0.500 B 0.920 B 0.095PIED 0.057 0.218 0.927 B 0.676 0.470 C 0.091 0.953 A 0.030 BPIEL 0.950 0.605 0.091 A B 0.213 0.953 C 0.500 0.057 0.927 B 1.040PIEN2 0.075 A 0.218 0.598 0.057 0.953 0.540 B 0.091 1.018 0.950 C B 0.030PIFL 0.370 0.598 0.215 0.057 B 0.780 C 0.953 0.155 0.075 B 0.598 B 0.953 0.215 0.950 A 0.091 0.057 C 0.370 B 0.927 0.215 A B 0.950 0.057 0.075 0.598 C 0.953 1.020 0.091 A B 0.075 0.950 C 0.215 0.598 0.927 B 0.057 B 0.010 0.075 B 0.598 0.950 0.215 0.091 B C 0.950 B 0.927 A 0.215 0.075 B 0.598 C B 0.927 0.075 0.091 0.950 0.727 C 0.215 B 0.091 0.927 0.598 B 0.075 B 0.213 C 0.091 0.927 0.215 B 0.991 B C 0.598 0.091 B C 0.598 0.089 0.215 0.927 B 0.215 0.598 B 0.091 C 0.215 0.598 C 0.598 B 0.215 C 0.215 0.598 C C 0.215 C exilis Speciestesota virginiana Codearboreum tomentosa den0borbonia rel1abies unc1breweriana cod1engelmannii rel2glauca lutzii unc2mariana cod2pungens rel3rubens unc3sitchensis cod3species rel4albicaulis aristata unc4arizonica cod4 attenuata balfouriana banksiana clausa contorta coulteri discolor echinata edulis elliotti engelmannii fl Genus Olneya Appendix D.—continued Ostrya Oxydendrum Paulownia Persea Picea Picea Picea Picea Picea Picea Picea Picea Picea Picea Pinus Pinus Pinus Pinus Pinus Pinus Pinus Pinus Pinus Pinus Pinus Pinus Pinus Pinus Pinus

35 continued PIGLPIJE 0.410PILA 1.038 0.370PILE 0.062 0.904 0.340PIMO 0.025 1.040 0.370 BPIMO3 0.500 0.038 0.953 BPIMU 0.987 0.350 0.953 0.057 B 0.078PINI 1.022 0.960 0.057 0.370 B 0.012PIPA 0.037 0.906 B 0.953 B 0.410PIPO 0.014 0.950 0.057 B B 0.540 0.953 0.991 0.950PIPU2 0.075 B 0.953 0.380 0.057 0.095 B 0.996 0.883 0.075PIRA 0.490 0.057 B 0.925 0.015 0.062 0.735 B B 0.950PIRE 1.038 B 0.011 0.370 0.054 B 0.927 0.075 BPIRI B 0.062 0.950 0.605 0.953 0.410 0.927 0.091 B BPISA 0.950 0.075 0.215 B 1.170 0.057 0.645 B 0.862 0.091 0.470 0.075 BPISE 1.007 0.055 0.214 0.517 0.052 B 0.370 C 0.927 0.953 B 0.987 B 0.020PIST2 0.213 B 0.598 0.953 0.091 B 0.510 0.057 C 0.078 B 0.927PIST 0.950 0.598 0.350 0.215 B 0.057 0.953 C 0.927 0.091 B 0.598 B 0.075PISY B 0.950 0.215 0.953 0.057 0.340 C 0.091 1.154 B 0.215 0.075PITA B 0.057 0.598 0.950 B 0.991 C 0.953 0.380 0.061 B BPIVI 0.950 C 0.215 0.075 0.095 B 0.057 0.598 0.990 B 0.470 0.927 B 0.075PLAQ 0.950 0.598 0.215 0.020 C 0.862 B 0.091 B 0.876 0.450 B 0.950 0.075PLOC 0.215 0.481 0.530 B 0.014 0.088 0.953 C A 0.075 0.927 B 0.605POAL 0.950 0.213 0.982 0.460 B C 0.057 0.927 B 0.091 0.215 BPOAN 0.075 0.980 B 0.129 0.982 0.598 0.370 C 0.091 0.927 B 0.020POBA 0.784 B 0.129 0.215 C 0.340 0.520 1.006 B 0.927 B 0.091 BPODE 0.012 1.006 0.214 0.132 0.310 0.950 A 0.091 0.598 B C 0.927 0.793 BPOFR 0.132 1.006 0.075 0.370 0.598 B 0.215 C B 0.091 0.830 B 0.201 0.793POGR 0.132 1.006 0.215 0.340 0.598 B B 0.080 0.732 C 0.201 0.793 BPOHE 0.598 0.132 B 0.360 1.006 0.215 B C 0.053 0.793 0.201 0.927 A 0.215 B 0.970 0.132 0.370 0.598 B 0.618 C 0.201 0.793 0.091 B 0.063 B 1.006 0.215 C 0.598 0.171 0.618 B 0.201 0.793 B 0.132 B 0.215 0.601 0.171 B 0.868 C 0.201 B 0.793 B 0.213 0.868 0.229 B 0.598 C B 0.793 0.201 B 0.525 0.229 0.868 0.215 C 0.201 B 0.793 0.223 0.525 B 0.229 0.868 B 0.201 C 0.223 B 0.613 0.229 C 0.868 B 0.613 0.227 B C 0.878 0.229 B 0.227 0.613 0.092 C 0.868 B 0.227 0.613 C 0.229 B 0.227 0.613 C B 0.613 0.227 C 0.227 0.613 C 0.227 C C Speciesglabra jeffreyi Codelambertiana leiophylla den0monophylla rel1monticola unc1muricata cod1nigra rel2palustris ponderosa unc2pungens cod2radiata rel3resinosa unc3rigida cod3sabiniana rel4serotina strobiformis unc4strobus cod4 sylvestris taeda virginiana aquatica occidentalis alba angustifolia balsamifera deltoides fremontii grandidentata heterophylla Genus Pinus Appendix D.—continued Pinus Pinus Pinus Pinus Pinus Pinus Pinus Pinus Pinus Pinus Pinus Pinus Pinus Pinus Pinus Pinus Pinus Pinus Pinus Pinus Planera Platanus Populus Populus Populus Populus Populus Populus Populus

36 continued POSAPOSP 0.370POTR 1.006 0.370PRSP 0.132 1.040 0.390PRAM 0.136 0.970 0.580 BPRNI 0.040 0.470 0.982 B 0.793PRPE 1.040 0.129 A 0.470 0.201 0.793PRSE 0.136 0.360 1.040 B 0.201 0.750PRSP2 B 1.040 0.470 0.136 B 0.050 0.793 0.470PRVI B 0.031 1.040 0.868 B 1.045 0.201 1.040PSME A 0.073 0.229 0.860 B 0.360 0.263 0.136QUAG 1.045 B 0.450 0.227 0.540 1.040 B B 1.011 0.263QUAL B 0.892 0.700 B 0.070 0.136 0.618 B 0.041 1.080QUAR 0.613 0.015 1.020 B 0.600 0.708 0.171 1.045 A B 0.059 0.227QUBI 0.613 0.051 B 0.700 1.012 0.192 B 0.263 0.708 B 0.227QUCH 0.613 1.045 B 1.020 0.032 C B 0.636 0.640 0.192 B 0.831 B 0.227 0.263QUCO 0.525 0.051 0.700 C 0.076 1.020 0.780 B 0.017 0.841 B 0.525 0.223QUDO 1.020 0.708 0.600 C 0.051 B 0.103 B 0.084 B 0.806 0.223QUDU 0.051 B 0.192 0.990 0.510 0.525 C B 0.708 B 0.841 0.035 BQUEL 0.052 0.525 1.020 0.223 0.600 C B 0.591 B 0.192 0.084 0.223 0.841QUEM 0.051 0.525 B 1.020 0.560 0.054 0.705 B C 0.841 0.525 0.084 B 0.223QUEN B 0.051 0.700 1.020 0.106 C B 0.591 0.084 B 0.223 0.904QUFA 1.020 0.051 0.700 B 0.525 C 0.705 0.078 B B 0.047 0.841 B 0.433 C 0.051 1.020 0.223 0.520 0.106 B 0.705QUFA2 0.084 B 0.841 0.213 0.591 B 0.051 1.020 0.705 B C 0.106 0.610QUGA B 0.084 0.841 0.216 B 0.398 0.051 C 0.106 0.781 B 1.020QUGA2 0.841 0.084 0.640 B 0.591 0.214 B C 0.078 0.705 0.051 B 0.640 BQUHY 0.084 1.020 0.841 0.216 B 0.591 0.106 C 0.705 1.020 BQUIL 0.051 0.084 B 0.700 0.841 0.591 B 0.216 C 0.106 0.051 0.705 BQUIM 1.020 0.084 0.216 0.591 B B 0.560 0.841 0.705 0.106 C B 0.051 B 0.216 0.591 0.560 1.020 0.084 B C 0.106 0.841 0.705 B 0.216 1.020 0.591 0.051 B 0.841 C 0.084 0.106 B 0.705 B 0.051 0.216 0.084 0.591 C B 0.841 0.106 B B 0.705 0.591 0.216 B C 0.084 B 0.841 0.106 B 0.216 0.705 0.591 C 0.841 0.084 B 0.705 0.106 0.216 B 0.591 C 0.084 0.106 B 0.705 0.216 B C 0.591 B 0.106 B 0.705 0.216 C 0.591 0.705 0.106 B 0.591 0.216 C 0.106 0.216 B 0.591 C B C 0.216 0.591 0.591 0.216 C 0.216 C C . var Speciessargentii species Codetremuloides species den0americana rel1nigra unc1pensylvanica cod1serotina rel2species virginiana unc2menziesii cod2agrifolia rel3alba unc3arizonica, grisea cod3bicolor rel4chrysolepis coccinea unc4douglasii cod4 durandii ellipsoidalis emoryi engelmannii falcata var. pagodaefolia gambelii garryana hypoleucoides ilicifolia imbricaria Genus Populus Appendix D.—continued Populus Populus Prosopis Prunus Prunus Prunus Prunus Prunus Prunus Pseudotsuga Quercus Quercus Quercus Quercus Quercus Quercus Quercus Quercus Quercus Quercus Quercus Quercus QuercusQuercus falcata Quercus Quercus Quercus Quercus

37 continued QUINQUKE 0.560QULA 0.510 1.020QULA2 1.020 0.051 0.520 0.560QULO 0.051 1.020 B 1.020QULY 0.051 0.640 B 0.051 0.841QUMA 1.020 0.570 B 0.841 0.084QUMA2 0.051 0.580 B 1.020 0.084 0.841 0.560QUMI 1.020 0.051 B B 0.841 0.084 1.020QUMU 0.051 B 0.600 0.084 B 0.705 0.051 0.841QUNI B 0.600 1.020 B 0.705 0.106 0.084 B 0.841QUNU B 1.020 0.051 0.106 0.560 0.705 0.841 0.084 B 0.051QUOB B 0.705 0.560 1.020 0.106 0.841 B 0.084 BQUPA 0.106 B 1.020 0.591 0.700 0.051 B 0.084 0.705 B 0.841 B 0.591QUPH 0.051 0.216 1.020 0.580 0.106 B 0.705 B B 0.841 0.084 0.216QUPR 0.051 0.591 1.020 0.560 0.705 0.106 B C 0.084 B 0.591 0.841 0.216 0.705QURU 0.051 1.020 B 0.570 0.106 C B 0.216 B 0.841 0.084 B 0.106 0.591QUSH 0.051 1.040 0.560 C B 0.705 B 0.084 0.841 0.216 C QUSP 0.591 0.079 1.031 0.560 B B 0.705 B 0.106 0.084 0.841 0.591 0.216 BQUST 0.044 1.020 0.106 0.580 C B 0.705 0.591 0.084 0.841 B 0.216 BQUVE 0.051 1.013 0.600 C 0.705 B 0.106 B 0.216 0.084 0.927QUVI B 0.021 0.591 C 1.020 0.560 0.106 0.705 B 0.086 0.859 B C 0.591QUWI B 0.216 0.051 1.020 0.106 0.705 B 0.800 B 0.074 0.841 0.216RONE B 0.051 0.591 0.700 0.106 0.705 C 1.020 B B 0.084 0.861 0.591 BROPS 0.216 C 0.660 1.020 0.106 0.051 0.547 B B 0.045 0.841 0.216 0.591SASP B 1.040 0.051 0.660 0.077 0.731 C B B 0.084 0.700 0.216 0.591SASE B 0.136 1.040 C 0.093 0.360 0.705 B B 0.072 0.216 0.591 0.841SAAL B 0.033 C 0.982 0.106 0.470 0.688 B B 0.841 0.216 0.084 0.591SESE B 0.129 C 0.982 0.079 0.420 0.705 B B 0.916 0.084 0.216 0.591SEGI 0.129 C B 1.040 0.340 0.106 0.858 B B 0.231 0.916 0.216 0.591 B C 0.030 0.994 0.118 0.340 B 0.705 B 0.055 0.793 0.216 0.686 B C 0.071 0.994 0.705 0.106 B B 0.201 0.793 0.213 0.591 B C 0.071 0.618 0.106 B 0.201 B 0.998 0.216 0.591 B C 0.171 0.618 B B 0.051 0.951 0.216 B 0.591 C 0.171 0.618 B 0.260 0.951 0.591 0.216 B C 0.171 0.618 B 0.260 0.525 0.216 B 0.171 C 0.853 B 0.223 0.525 B C 0.077 0.902 B 0.223 0.525 C 0.146 0.902 B 0.223 0.525 C 0.146 B 0.223 0.525 C B 0.223 0.605 C 0.218 0.605 C 0.218 C C Speciesincana kelloggii Codelaevis laurifolia den0lobata rel1lyrata unc1macrocarpa cod1marilandica rel2michauxii muehlenbergii unc2nigra cod2nuttalli rel3oblongifolia unc3palustris cod3phellos rel4prinus rubra unc4shumardii cod4 species stellata velutina virginiana wislizeni neomexicana pseudoacacia species sebiferum albidum sempervirens giganteum Genus Quercus Appendix D.—continued Quercus Quercus Quercus Quercus Quercus Quercus Quercus Quercus Quercus Quercus Quercus Quercus Quercus Quercus Quercus Quercus Quercus Quercus Quercus Quercus Quercus Quercus Robinia Robinia Salix Sapium Sassafras Sequoia Sequoiadendron

38 continued SOAMTASP 0.420 0.982 0.400TADI 0.129 0.982TABR 0.420 0.129 BTHOC 0.600 0.994 BTHPL 0.793 0.290 0.994 0.071TIAM 0.201 1.040 0.071 0.793 0.310 BTIHE 0.039 0.201 1.040 B 0.320 BTISP 0.951 0.074 1.040 B 0.320 B 0.618 0.951 0.260TOCA 0.026 1.040 0.320 B 0.171 0.960 0.260 0.618TSCA 0.340 B 0.136 1.040 B 0.163 0.171 0.960 BTSHE B 0.994 0.400 0.136 0.902 B 0.326TSME 1.080 B 0.071 B 1.035 0.440 0.525 0.902 0.146 B 0.046TSSP 1.080 0.060 B 0.900 0.223 0.420 1.330 0.146 B 0.525 B 0.272ULAL 1.080 0.030 0.953 B 0.075 0.380 0.223 B 1.064 C B 0.951ULAM 0.272 0.052 0.605 0.953 B 0.570 0.058 A 0.848 B 0.260 C 0.882ULCR 0.605 0.218 0.052 0.460 0.982 B B 0.076 0.848 0.241 B 0.830 0.655ULPU 0.218 B 0.982 0.129 0.570 C B 0.225 0.848 0.040 0.882 B 0.213ULRU 0.129 B 0.656 0.982 0.460 C 0.902 B 0.225 0.241 0.882 BULSE 0.213 A 0.129 0.525 C 0.982 0.480 0.146 B 1.081 0.241 0.793 BULSP B 0.223 0.129 0.982 0.525 0.570 C 0.087 0.661 B B 0.793 0.201ULTH 0.129 B 0.223 0.982 0.525 0.500 0.050 0.906 C B 0.201 B 0.793UMCA 0.605 0.129 B 0.223 0.982 0.079 0.570 B 0.906 C A 0.201 0.793 0.218VAAR B 0.604 0.510 0.129 0.982 0.079 B 0.618 C B 0.201 0.793 0.218 0.380 B 0.982 0.129 C 0.470 0.618 0.171 B 0.201 B 0.793 0.050 0.604 0.129 B 0.982 C 0.171 0.618 B 0.201 B 0.793 0.218 B 0.604 0.129 A 0.171 B 0.618 B 0.201 0.793 0.218 B 0.525 C 0.171 0.618 B B 0.793 0.201 0.525 0.223 B C 0.171 0.618 0.201 B 0.793 0.223 0.525 B C 0.171 0.618 B 0.201 0.223 B 0.525 C 0.171 0.618 B 0.223 0.525 B C 0.618 0.171 B 0.223 0.525 0.171 C 0.618 B 0.223 0.525 C 0.171 B 0.223 0.525 C B 0.525 0.223 C 0.223 0.525 C 0.223 C C Speciesamericana species Codenutans den0brevifolia rel1occidentalis plicata unc1americana cod1heterophylla rel2species unc2californica cod2canadensis rel3heterophylla mertensiana unc3species cod3alata rel4americana unc4crassifolia cod4 pumila rubra serotina species thomasii californica arboreum Genus Sorbus Appendix D.—continued Tamarix TaxodiumTaxus distichum var. Thuja Thuja Tilia Tilia Tilia Torreya Tsuga Tsuga Tsuga Tsuga Ulmus Ulmus Ulmus Ulmus Ulmus Ulmus Ulmus Ulmus Umbellularia Vaccinium

39 3 BC Genera sampled Species and genus not sampled BC Genera sampled Species and genus not sampled BC Genera sampled Species and genus not sampled BC Genera sampled Species and genus not sampled nition Code/Units Method fi Appendix D.—continued Standing Dead Tree Relative Density Predictions Metadata (Appendix D) Fieldname De rel1unc1cod1 relative density decay class 1 uncertainty of rel1 dimensionless uncertaintly code for rel1 A dimensionless Species sampled genusspeciescode genus species den0 genus-species code green densityrel2 g/cm unc2cod2 relative density decay class 2 uncertainty of rel2 dimensionless uncertaintly code for rel2rel3unc3 Acod3 dimensionless relative density decay class 3 uncertainty of rel3 dimensionless uncertaintly code for rel3rel4 Species sampled unc4 Acod4 dimensionless relative density decay class 4 uncertainty of rel4 dimensionless uncertaintly code for rel4 Species sampled A dimensionless Species sampled

40 Harmon, Mark E.; Woodall, Christopher W.; Fasth, Becky; Sexton, Jay; Yatkov, Misha. 2011. Differences between standing and downed dead tree wood density reduction factors: A comparison across decay classes and tree species. Res. Pap. NRS-15. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station. 40 p.

Woody detritus or dead wood is an important part of forest ecosystems and has become a routine facet of forest monitoring and inventory. Biomass and carbon estimates of dead wood depend on knowledge of species- and decay class-specifi c density or density reduction factors. While some progress has been made in determining these parameters for dead and downed trees (DD), there are very few estimates of these key parameters for standing dead trees (SD). We evaluated indicators of decay to relate subjective SD and DD decay classifi cations then compared SD and DD density and density reduction factors by decay class for a total of 19 tree species at nine sites in the United States and Russia. Results indicate that SD density declined with decay class for all examined species. By applying these results, a new set of SD density reduction factors was developed for 260 species inventoried by the U.S. Forest Service’s Forest Inventory and Analysis program in forests of the United States.

KEY WORDS: standing dead trees, downed dead trees, carbon, biomass, decay reduction factor

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